Biomass production and nutrient uptake from cover crops in Brazilian agriculture

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In Brazil, however, their use remains limited due to technical barriers and a lack of accessible data. In this systematic literature review, we extensively searched for data on biomass production and nutrient uptake of the primary cover crop families used in Brazilian agriculture. We aimed to map available research, identify knowledge gaps, and offer data to inform cover crop adoption. We retrieved over 1,300 articles from three databases, selecting 126 studies that yielded more than 1,000 observations of cover crops grown as single species or in mixtures. In total, 66 species and 58 multispecies mixes from seven botanical families were represented. Cover crop biomass production ranged from 0.33 to 17.9 Mg ha⁻¹, with an average carbon content of 44.8%, and C inputs varying from 489 to 2,898 kg ha⁻¹, decreasing from Poaceae to Brassicaceae . Nitrogen uptake ranged from 17 to 119 kg ha⁻¹, with the highest values in Fabaceae and similar uptake in Brassicaceae and Poaceae mixtures. Phosphorus and potassium uptake varied across species, with Poaceae showing the highest phosphorus and potassium, while mixtures had varied nutrient content, with limited data on phosphorus and potassium. Overall, these findings highlight the variability in nutrient uptake among different cover crop species and mixtures, providing valuable insights for optimizing cover crop selection to enhance soil health and nutrient management. green manure soil health regenerative agriculture functional biodiversity nutrient cycling Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Conventional agricultural systems simplify natural ecosystems to maximize the provision of one ecosystem service (i.e., provision of food, feed, fiber, and fuel) to the detriment of the provision of other services (Power, 2010 ; Pereira et al., 2018 ; Hasan et al., 2020 ). Conservation agriculture practices were developed in the last decades as a framework of management strategies to deter, mitigate, and reverse soil organic carbon (SOC) stock depletion (Poeplau and Don, 2015 ; He et al., 2025; Wu et al., 2025 ) and other ecosystem services in agroecosystems (Lal, 2015 ; Jian et al., 2020 ; Yang et al., 2024 ). It is based on new and age-old nature-based solutions (NBS) implemented, to various degrees of adoption, in modern agroecosystems, such as organic fertilization, biological control, integrated systems, no-till, and plant diversification (Ahmad et al., 2021 ; Yost et al., 2022 ; Prairie et al., 2023). Diversification aims to reverse the loss of soil multifunctionality in agroecosystems. It does so by incorporating functional biodiversity through specific groups of plants. This approach helps restore complex biotic interactions. Consequently, these interactions enhance the delivery of multiple ecosystem services in agroecosystems (Daryanto et al., 2018 ; Yousefi et al., 2024 ). Plant diversification can be implemented in agricultural systems by the incorporation of cover crops. Cover crop use is an ancient management practice with multiple potential benefits to soil health, crop yields, and the environment (Koudahe et al., 2022 ; Yang et al., 2024 ; Yin et al., 2025 ). Cover crops directly or indirectly support the supply of several soil functions and soil-related ecosystem services such as nutrient cycling and provision, water filtering and flow regulation, climate regulation, erosion control, and pest population control (Van Eerd et al., 2023 ; Qiu et al., 2024 ; Schön et al, 2024 ). These benefits are related to the capacity of cover crops to increase SOC stocks in agroecosystems (Schlesinger, 2022 ). In tropical climates, cover cropping can increase SOC by 7.2% on average (Jian et al., 2020 ). In the Brazilian context, the edaphoclimatic conditions and management practices from north to south of the country provide a wide range of options for cultivating cover crops throughout the agricultural year (Cherubin et al., 2024 ). In tropical agricultural systems, cover crops are often classified into functional groups of families based on the specific ecosystem services they can provide to agroecosystems. For example, tropical grasses can accumulate high biomass in a short period, resulting in biomass left in the field covering the soil surface to prevent erosion and nutrient loss (e.g., Baptistela et al., 2020; Souza et al., 2025 ). These species also possess vigorous root systems capable of scavenging water and nutrients at great soil depths and improving subsequent cash crop production (Costa et al., 2021 ). However, the thick layer of residue can hinder planting operations, and depending on the litter stoichiometry, it can immobilize nutrients, limiting their availability throughout the cash crop growing season. Although legume cover crops typically produce less biomass, they fix atmospheric nitrogen and generate residues with lower C:N ratios, promoting nutrient mineralization and availability during the subsequent cash crop growing season (Vendig et al., 2023 ; Peng et al., 2024 ). While cover crops provide numerous beneficial ecosystem services (e.g., Blanco-Canqui et al., 2015 ; Daryanto et al., 2018 ; Yousefi et al., 2024 ), they may also negatively impact other services or crop management practices (e.g., Glaze-Corcoran et al., 2023 ; Rouge et al., 2023 ). The concurrent growth of cover crops and cash crops in the same space can result in both plants competing for the same resources, even in succession systems (Rouge et al., 2023 ). Cover crops are often recommended as a strategy to supply nutrients for the subsequent crop during the litter decomposition; however, the timing of litter decomposition and nutrient release often does not align with the crop’s demand (Nevins et al., 2020 ). For instance, grasses with a high C:N ratio can immobilize nitrogen during decomposition, delaying its availability to the following crop and potentially causing nitrogen deficiency and yield reductions (Deines et al., 2023 ). Thus, a better understanding of biomass production and nutrient accumulation in the biomass of different groups of cover crops is urgent to provide adequate cover cropping recommendations and avoid or mitigate negative trade-offs. This data can also be used to refine fertilization recommendations, determine the amount of C added to the system, and develop adequate C models or C footprint calculators. Many challenges regarding cover cropping are still posed to Brazilian researchers, consultants, and farmers. The technical constraints include seed availability, high logistical costs, and access to specialized equipment for cover crop termination. In addition, the scarcity of organized data on biomass production and nutrient extraction by cover crop species across different edaphoclimatic conditions limits the ability of researchers to design more accurate recommendations. Finally, the use of cover crop mixes adds yet another layer of complexity to the questions that remain open for investigation. To fill the basic knowledge gap, in this study, we present an extensive systematic literature review of the available experimental data for cover crops in Brazil, including biomass production and nutrient extraction. The data will assist farmers and consultants in the decision-making process of choosing adequate cover crop species or mixtures for increased soil health in Brazilian farming systems, as well as researchers and policymakers in developing region-specific guidelines, advancing soil management research, and shaping sustainable agricultural practices. Material and methods Search and dataset preparation The search for articles was restricted by inclusion and exclusion criteria established to find the relevant literature for the systematic review. The inclusion criteria were: i) the publication must be a peer-reviewed article published in a scientific journal; ii) the paper must include cover crop species; iii) the paper must include biomass production, and the uptake by the cover crop of at least one of the following elements: carbon, nitrogen, phosphorus, potassium, calcium, and magnesium; iv) the paper must include the experiment duration with planting and termination dates; v) the studies must be conducted in the field; vi) the experiment must be conducted in Brazil, and vii) the paper must contain the exact or approximate location of the experiment. The exclusion criteria used were: i) studies conducted in semi-perennial systems (e.g., sugarcane), orchards (e.g., oranges, coffee), or vegetable gardens; ii) studies with non-conventional cover crop species (e.g., Calotropis procera ); iii) results reported in non-compatible measurement units (e.g. g/plant, g/pot); and iv) results presented as the average of two or more cropping years. The search for articles in the Scopus and Web of Science databases was done using keywords in English. In the Scielo database, the search was in English and Portuguese. The intent of using two languages was to access the highest number of published literature in Brazilian databases, including articles published in academic journals not available in English. The search considered terms mentioned in each record's "title, abstract, and keywords" (Topic Field). The search query in English used was: ("cover crop*" OR "green manur*") AND ("biomass" OR "dry matter"). The search query in Portuguese for the Scielo database was: ("planta* de cobertura" OR "adub* verde") AND ("biomassa" OR "matéria seca"). The search was restricted to studies from Brazil using the “country/region” filtering tool available on the platforms. The search on the three databases occurred on 21 May 2022. Initially, a total of 1372 papers were retrieved by the search (Fig. 1 ). All the retrieved articles were organized using reference management software and sorted to remove duplicates. Thus, the number of papers in the dataset was reduced to 840 (Fig. 1 ). Each unique paper retrieved from the databases was manually sorted by the inclusion/exclusion criteria pre-established before the search. The papers matching all criteria were selected to build the raw dataset used for statistical analysis. The sorting reduced the number of articles from the original 840 to 126 selected articles (Fig. 1 ). Subset data within the raw data frame From the 126 articles selected for the raw database, data were extracted on biomass production, scientific names of the cover crop species, geographic coordinates, planting season, experiment duration, soil texture, and soil classification. Data reported in non-standard units were converted to the standard measurement (i.e., kg ha⁻¹). Observations of biomass production below 300 kg ha − 1 or exceeding 18 Mg ha − 1 were also removed as they were considered outliers for any cover crop within a field study with a growing period ranging from 30 to 200 days, the latter of which would allow for at least 156 days for one or more cash crop harvest (Fig. 1 ). The data was organized into a spreadsheet, totaling 1165 individual observations. Using this database, the data were subdivided by grouping each species of cover crop into its respective taxonomic families. After the data filtering, we obtained 758 observations of Fabaceae , Poaceae , Brassicaceae , Asteraceae , and other less common families in monoculture, as well as 199 observations of multispecies mixtures of Fabaceae and Poaceae , two genera or species of Poaceae , Brassicaceae , and Poaceae , and other less common mixtures. Data analysis The frequency of observations of biomass, the N, P, and K uptake, planting months, and the growing period were organized in histograms by taxonomic family. An exploratory statistical analysis was performed to obtain each variable's mean, median, and standard deviation using standard statistical libraries from the R software. Results We sought to obtain an overview of the biomass production, C inputs, and N, P, and K uptake by cover crops from botanical families commonly used in Brazilian agriculture. The main region represented in studies was Central Brazil, with over 72% of total observations and 68% of total unique studies. The next region represented was Southern Brazil, below the Tropic of Capricorn, with 21% of observations and 22% of unique studies (Fig. 2 a). In terms of climate, 53% of the study sites are in the humid subtropical regions, 46% in tropical regions, and less than 1% in dry regions (Fig. 2 c). Regarding soil classification, 69% of the sites are on Ferralsols, 17% on Acrisols, 9% on Cambisols, and 5% on other soil types, including Arenosol, Fluvisol, Lixisol, Nitisol, and Plinthosol (Fig. 2 d). Sixty-six species of cover crops and 58 multispecies mixes were retrieved from the 119 articles included in this systematic review. Overall, they belong to seven botanical families, i.e., Poaceae , Fabaceae , Asteraceae , Brassicaceae , Amaranthaceae , Polygonaceae , and Pedaliaceae . The Amaranthaceae , Polygonaceae , and Pedaliaceae families had fewer than 10 observations each and were grouped as “other families” for the data analysis. The biomass production ranged from 0.33 to 17.9 Mg ha − 1 , and the average C content was 44.8% of the total biomass. The C inputs to the system ranged between 489 and 2,898 kg ha − 1 , decreasing from Poaceae (3076 kg ha − 1 ), Asteraceae (2781 kg ha − 1 ), Fabaceae (2393 kg ha − 1 ), to Brassicaceae (1727 kg ha − 1 ) when planted as a single cover crop species. The general N uptake of single cover crop species ranged from 17 to 119 kg ha − 1 . The highest mean values were found for the Fabaceae (121 kg ha − 1 ) family, followed by Poaceae (107 kg ha − 1 ), Brassicaceae (80 kg ha − 1 ), and Asteraceae (51 kg ha − 1 ) (Fig. 3 ). The mean P uptake of the cover crops ranged from 4.8 to 24 kg ha − 1 from Poaceae (24.2 kg ha − 1 ), Asteraceae (17.5 kg ha − 1 ), Fabaceae (15.3 kg ha − 1 ), to Brassicaceae (4.8 kg ha − 1 ) (Fig. 3 ). The K uptake ranged from 33 to 116 kg ha − 1 , decreasing from Poaceae (132 kg ha − 1 ), Asteraceae (105 kg ha − 1 ) and Fabaceae (95 kg ha − 1 ) to Brassicaceae (51 kg ha − 1 ) when planted as single species (Fig. 3 ). For multispecies mixtures, the C inputs varied from 486 to 2,666 kg ha − 1 , decreasing from the Fabaceae with Poaceae mix (2860 kg ha − 1 ), to Brassicaceae and Poaceae mixes (2461 kg ha − 1 ), and to the mix of two Poaceae (1552 kg ha − 1 ), which resulted in the lowest C inputs. Multispecies mixtures of different families resulted in the highest C inputs among the mixtures. The cover crop mixtures with plants of the Brassicaceae and Poaceae (149 kg ha − 1 ) families, and the Fabaceae and Poaceae mixtures (127 kg ha − 1 ) had similar N uptake. However, data on Brassicaceae and Poaceae mixtures were significantly scarcer than other mixtures ( n = 9 ). Data on P and K content were reported for the Fabaceae and Poaceae (17 and 107 kg ha − 1 ) mix, and the mix of two or more Poaceae (2 and 31 kg ha − 1 ) (Fig. 3 ). The most prevalent cover crop mixtures included plants from the Fabaceae and Poaceae families, accounting for 51% of all observations, followed by mixtures with two or more species of Poaceae with 31% of observations. To a smaller extent, Brassicaceae was also represented in studies with cover crop mixtures, with 27 observations of Brassicaceae species being used in mixtures with plants from the Poaceae family. Fabaceae A total of 82 studies resulted in 415 observations of biomass production of Fabaceae species within the established constraints. A total of 21 genera of plants belonging to the Fabaceae family were found in the dataset and can be consulted in the supplementary material ( Table S1 ). Biomass production values for Fabaceae range from 0.33 to 17.85 Mg ha − 1 with a mean, median, and standard deviation of 5.34, 4.38, and 3.90 Mg ha − 1 , respectively (Fig. 4 a - blue line). The median extraction of N, P, and K is 105, 9, and 60 kg ha − 1 . Fabaceae has a median growth period between planting and desiccation of 104 days (Fig. 4 f), with planting operations occurring more frequently between either February and June or in November, although the data show planting in all months of the year (Fig. 4 e). Species ideally planted in the summer months, like Cajanus cajan (5.9 Mg ha⁻¹) and Crotalaria juncea (6.7 Mg ha⁻¹) had a median biomass production of 5.0 Mg ha⁻¹, while the winter cover crops, such as Lupinus spp. (4.1 Mg ha⁻¹) and Vicia spp. (2.8 Mg ha⁻¹), had a median of 2.59 Mg ha⁻¹ ( Table S1 ). In the subtropical region of Brazil, below latitude 23°S, the median biomass production was 5.7 Mg ha⁻¹ ( n = 54) in the summer and 2.88 Mg ha⁻¹ ( n = 15) in the winter. Poaceae A total of 84 studies resulted in 422 observations of biomass production for the Poaceae family. A total of 14 genera of plants belonging to the Poaceae family were found in the dataset and can be consulted in the supplementary material ( Table S1 ). Biomass production of Poaceae ranged between 0.35 and 17.73 Mg ha − 1 with a mean, median, and standard deviation of 6.86, 6.21, and 4.13 Mg ha − 1 , respectively. The median extractions of N, P, and K from Poaceae are 96, 14, and 99 kg ha − 1 over a mean growth period between planting and desiccation of 109 days (Fig. 4 f - green line), with most plantings occurring between February and May (Fig. 4 e). The biomass production reported for the Poaceae family spanned from 0.35 to 17.73 Mg ha⁻¹, and a median value of 6.21 Mg ha⁻¹. In the tropical region, the three main grasses planted were Cenchrus spp., with median biomass production of 5.3 Mg ha⁻¹; Sorghum bicolor with 3.8 Mg ha⁻¹; and Urochloa ruziziensis with 9 Mg ha⁻¹. The most frequently used species below latitude 23°S were Avena spp. and Cenchrus spp., followed by Urochloa spp.. The median biomass production of these species was 4 Mg ha⁻¹, 8.6 Mg ha⁻¹, and 1.2 Mg ha⁻¹. Brassicaceae A total of 19 studies resulted in 42 observations of biomass production of Brassicaceae . We found 2 genera of Brassicaceae used in the field, namely Raphanus and Crambe ( Table S1 ). Biomass production of Brassicaceae ranges from 0.73 to 8.30 Mg ha − 1 with a mean, median, and standard deviation of 3.85, 3.59, and 1.78 Mg ha − 1 , respectively. The median extractions of N, P, and K from Brassicaceae are 66, 6, and 60 kg ha − 1 over a mean growth period between planting and desiccation of 109 days (Fig. 4 f - purple line), with the most frequent plantings occurring between March and May (Fig. 4 e). No studies were observed for planting dates in January, August, September, or December (Fig. 4 e). Brassicaceae exhibited a biomass production ranging from 0.73 to 8.30 Mg ha⁻¹, with a median of 3.59 Mg ha⁻¹. The main species used was Raphanus sativus , commonly planted as a winter cover crop in the subtropical region. Median P and K uptakes for the family were 6 kg ha⁻¹ and 53 kg ha⁻¹. The mean C:N ratio was consistent with the ratio for a localized long-term study with R. sativus , around 22 (Balota et al., 2014 ). Asteraceae A total of 6 studies resulted in 14 observations of biomass production for the Asteraceae family. For the two genera of Asteraceae used in the field, biomass data were compiled for Helianthus and Thitonia ( Table S1 ). Biomass production of Asteraceae ranges from 0.90 to 15.20 Mg ha − 1 with a mean, median, and standard deviation of 6.21, 6.47, and 3.41 Mg ha − 1 , respectively. The median extractions of N, P, and K from Asteraceae are 65, 4, and 116 kg ha − 1, respectively, over a mean growth period between planting and desiccation of 112 days (Fig. 4 f - red line), with the most frequent plantings occurring between February and March. No studies were observed for plantings in January, April, and between July and December (Fig. 4 e). The main species used was Helianthus annuus , also a cash crop planted for vegetable oil production. Median P and K uptakes for the family were 1.7 kg ha⁻¹ and 105 kg ha⁻¹. The median C:N ratio for the family was 39. Cover crop mixtures The most used mixtures in the selected studies were combinations of Fabaceae and Poaceae species, with 32 studies resulting in 256 observations. The biomass production of the mixtures of Fabaceae and Poaceae species ranged between 0.63 and 17.00 Mg ha − 1 with the mean, median, and standard deviation of 6.38, 5.71, and 3.45 Mg ha − 1 , respectively (Fig. 5 a - blue line). The median of the N, P, and K extractions were 104, 11, and 80 kg ha − 1, with an average planting period between planting and termination of 107 days, and the most frequent planting months were between March, May, and November (Fig. 5 e,f). The most common mixture for the subtropical region of Brazil was Avena strigosa with Vicia sativa , and Cajanus cajan with Urochloa ruziziensis for the tropical region. Medians of biomass production for the subtropical and tropical regions were 4.4 Mg ha⁻¹ and 6.6 Mg ha⁻¹. We found similar P uptakes for both regions, with 14.2 Mg ha⁻¹ and 13.6 Mg ha⁻¹ for the subtropical and tropical regions. The median K uptakes were also similar, 121 Mg ha⁻¹ and 103 Mg ha⁻¹ for the subtropical and tropical regions. The C:N ratio for the subtropical region with Fabaceae and Poaceae species was 22.8, while tropical mixtures had a higher C:N ratio of 40. Seven studies with a total of 27 observations presented data for mixtures of Brassicaceae and Poaceae species. We identified 5 different combinations of species of the two families, all of them with Raphanus sativus . The biomass production of the mixtures of Brassicaceae and Poaceae ranged between 0.64 and 10.40 Mg ha − 1 with the mean, median, and standard deviation of 5.49, 5.22, and 2.74 Mg ha − 1 , respectively. The median of the N extraction was 106 kg ha − 1 with the average growing period between planting and termination of 107 days, and the most frequent planting months were between March and May (Fig. 5 e,f - purple line). No P and K concentrations were reported for mixtures of Brassicaceae and Poaceae in the studies retrieved and selected in this systematic review. Mixtures of two Poaceae species were identified in 8 studies with 67 total observations. We identified 11 different combinations of species. The biomass production of these mixtures ranged between 0.33 and 16.40 Mg ha − 1 with the mean, median, and standard deviation of 3.46, 1.09 e 4.10 Mg ha − 1 , respectively. The medians of the N, P, and K extraction were 17, 1, and 33 kg ha − 1 with the growing period between planting and termination of 120 days, and the planting months were mostly between March and April (Fig. 5 e,f - green line). Discussion Biomass production and nutrients uptake This systematic literature review provides insights into biomass production and nutrient extraction of cover crops in Brazilian agroecosystems, linking family traits to ecosystem services. Fabaceae and Poaceae families dominated the studies, accounting for over 92% of the observations. Species from the genera Urochloa , Crotalaria , and Cenchrus represented 40% of these occurrences, largely due to their proven effectiveness in improving soil health and nutrient cycling, as well as their hardiness and strong adaptation to Brazilian climatic and soil conditions. As the highest biomass-producing family of cover crops, species of Poaceae achieved a median production of 6.21 Mg ha⁻¹. This result supports earlier reports that tropical grasses excel in biomass production thanks to efficient nutrient uptake and deep, extensive roots, traits well documented for Urochloa ruziziensis (Baptistella et al., 2020 ). Fabaceae displayed moderately high biomass (median 4.38 Mg ha⁻¹), underscoring their dual contribution of substantial biomass alongside N fixation capabilities, particularly notable in tropical legumes such as Crotalaria juncea and Cajanus cajan (Berriel and Perdomo, 2023 ). The intermediate productivity observed in Brassicaceae (median 3.59 Mg ha⁻¹) aligns with their recognized rapid growth but limited overall biomass compared to grasses (Elhakeem et al., 2023 ). Interestingly, Asteraceae also presented high biomass yields (median 6.47 Mg ha⁻¹), higher than most species of the Poaceae family and indicating variability possibly influenced by species selection, climate conditions, and management strategies across studies (Freitas et al., 2019). Tropical grasses like Urochloa spp. are preferred in the Brazilian savannah for rapid growth, drought tolerance, and high biomass yield, whereas in subtropical areas, grasses like ryegrass dominate due to cold tolerance and forage potential (Pavinato et al., 2014 ; Baptistella et al., 2020 ). However, biomass yields of tropical grasses can be lower in subtropical conditions, as shown by median biomass production of U. brizantha (6.5 Mg ha¹ in tropical; 3.4 Mg ha¹ in subtropical), emphasizing the necessity for selecting species suitable for local edaphoclimatic conditions to optimize agroecosystem benefits. Furthermore, cover crops classified as Fabaceae , particularly species of Crotalaria , are strategically used by farmers to control plant-parasitic nematodes sustainably, an issue increasingly critical under climate change conditions, which may enhance nematode survival rates (Lopes et al., 2019 ; Khanal and Land, 2023 ; Souza et al., 2024 ). Moreover, Fabaceae exhibited the highest N content and lowest C:N ratio (median C:N ratio of 16.9), consistent with studies in Brazilian savannah agroecosystems (Carvalho et al., 2013 ). Although individual species can provide ecosystem services, the combination of more than one species can balance different services, such as mixing grasses with legumes for balancing the high biomass production of the first and the residue quality (lower C:N) of the second (Plumhoff et al., 2022). In this study, the use of multispecies mixtures followed similar single-species trends and provided complementary functions and broadened ecosystem services. Fabaceae and Poaceae combinations made up 57% of observations, followed by mixtures with multiple Poaceae species and Poaceae cover crops intercropped with cash crops. The most common use of mixtures of Poaceae and Fabaceae species can be explained by plant complementary traits and functions (Schipanski et al., 2014 ). The mixtures incorporate both the N-fixing capabilities of Fabaceae species with the high biomass potential of Poaceae . The goal of using these mixtures then is to provide greater soil cover with lower N immobilization for the subsequent cash crop, and higher C accrual. This approach also aligns with the understanding that effective soil C sequestration depends on balancing C inputs with N availability, as soil organic matter formation and stabilization require proportionally more N than plant biomass (Cotrufo et al., 2015 ). Moreover, mixtures of Poaceae and Fabaceae have been shown to improve soil health and reduce yield variability over time, promoting greater resilience to climate-related stresses (Souza et al., 2025 ). Yet the use of mixtures demands greater management expertise from farmers. Per our results, biomass production for Fabaceae and Poaceae mixtures varied from 0.63 Mg ha⁻¹ to 17 Mg ha⁻¹, with an overall median of 5.7 Mg ha⁻¹. The range is due to differences in climatic conditions, planting seasons, and species of choice, the same as for single species. In addition, initial seeding proportions of mixtures, when available in the consulted literature, were highly variable and could have influenced the evaluated variables. Our results also revealed that mixtures used in the subtropical region, usually planted in winter, allow for the preponderance of Fabaceae plants with lower C:N ratios, while mixtures used in the tropics have C:N ratios much more similar to general Poaceae , indicating a preponderance of these plants in the mixture. In Brazil, intercropping cover crops with cash crops (Santa Fé System) using Urochloa grasses (mainly U. ruziziensis and U. brizantha ) is common, primarily to protect soil from erosive tropical rainfall events. The mean biomass produced by Urochloa grasses is approximately 5.7 Mg ha⁻¹, varying according to the cash crop due to competition for light and nutrients. Biomass production is higher with soybean (5.3 Mg ha⁻¹) than with maize (4.19 Mg ha⁻¹), while intercropping with common beans achieves 7.9 Mg ha⁻¹, although studies with beans are limited. While intercropping usually does not significantly impact maize yield (Brito et al., 2023 ; Souza et al., 2024 ), soybean yields can be negatively affected if not properly managed, necessitating selective herbicide use (Saraiva et al., 2013 ). Additionally, intercropped Urochloa grasses facilitate integrated crop-livestock systems by providing forage during dry winters, enhancing livestock productivity, and diversifying farmers' income (Souza et al., 2024 ). Beyond biomass production, the same families differed markedly in their ability to capture and recycle key nutrients. Fabaceae fix abundant N (median 105 kg ha⁻¹) yet yield moderate biomass, whereas Poaceae generate the greatest biomass and recover similar N (96 kg ha⁻¹) but much more K (99 kg ha⁻¹), a reflection of deep roots and rapid growth (Passot et al., 2016). Brassicaceae show swift early uptake, removing 66 kg N and 60 kg K ha⁻¹ within about 109 days while alleviating compaction via tap-roots (Tribouillois et al., 2015). Their rapid growth makes them excellent plants to cover the short fallow periods between harvests in the Brazilian crop succession systems common in the subtropical South and irrigated areas of western Bahia (Altieri et al., 2011 ). Asteraceae , dominated by sunflower in this review, accumulate high K (116 kg ha⁻¹) but lower N and P, and the high C:N ratio (> 40) suggests slower mineralization and longer soil cover. Mixtures such as millet and crotalaria synergize N fixation with K capture, boosting total nutrient return and subsequent crop yields over monocultures (Silva et al., 2010; Souza et al., 2025 ). Phosphorus and K uptake by cover crops significantly varied between subtropical and tropical regions, reflecting regional differences in species selection and biomass accumulation. Implications When biomass and nutrient profiles are viewed together, clear agronomic patterns emerge that inform agronomic decisions for optimizing agricultural practices and sustainable soil management strategies. Biomass production varies widely (0.33 to 17.9 Mg ha⁻¹), with average C inputs by families ranging from 489 to 2,898 kg ha⁻¹. This information is especially relevant for modeling soil carbon stocks and estimating the carbon footprint of agricultural systems (Jordon et al., 2022; McClelland et al., 2021; Prairie et al., 2023). Additionally, cover crops play a critical role in climate adaptation by leaving a protective mulch layer on the soil surface, especially when species with high C:N ratios are used. This mulch moderates diurnal temperature fluctuations and reduces evaporation, creating a more stable microclimate that enhances the resilience of subsequent cash crops to heat and drought stress (Souza et al., 2025 ). In particular, Poaceae species such as Urochloa ruziziensis and Sorghum bicolor are known for their high biomass production (median of 6.21 Mg ha⁻¹) and high C:N ratios, which contribute to slower decomposition and prolonged soil cover (Souza et al., 2025 ; Carvalho et al., 2013 ). With carbon inputs ranging from 489 to 2,898 kg C ha⁻¹, many cover crop species, especially in mixtures, provide enough biomass to match or substantially reduce the typical C deficits observed in degraded tropical soils which can vary from 100 kg C ha − 1 y − 1 to 3150 kg C ha − 1 y − 1 (Medeiros et al., 2021; Oliveira et al., 2022). These inputs are particularly important because they help replenish particulate organic carbon (POC), the soil carbon fraction most vulnerable to degradation under intensive agriculture. As POC plays a key role in soil structure, nutrient retention, and microbial activity, its restoration through high-biomass regenerative practices is essential for rebuilding soil health and long-term productivity (Prairie et al., 2023). Furthermore, biological nitrogen fixation by Fabaceae can reduce the need for external N inputs (Schipanski et al., 2014 ; Skinner et al., 2014 ). This reduction is particularly relevant for climate mitigation, as the production and application of synthetic nitrogen fertilizers are major sources of greenhouse gas emissions, especially nitrous oxide (N₂O), which has a global warming potential 265 times greater than CO₂ (IPCC, 2021). These results also provide valuable insights for designing multi-species cover crop mixtures in Brazil. By quantifying family-specific nutrient uptake and identifying a median growth window of 104 to 112 days, the findings support mixtures that combine N-fixing Fabaceae with high-biomass Poaceae , supplemented by smaller proportions of Brassicaceae for pest suppression. These combinations can be timed to regional climatic windows so that sowing dates maximize biomass and ensure that residue decomposition and N release coincide with cash‑crop demand. The data highlights the strong performance of Poaceae in nutrient uptake, particularly phosphorus (24 kg ha⁻¹) and potassium (131 kg ha⁻¹), underscoring their key role in enhancing nutrient cycling. When combined with other functional groups, such as Fabaceae and Brassicaceae , these grasses contribute to more balanced cover crop mixtures that support efficient nutrient use and reduce the need for supplementary chemical fertilization (Souza et al., 2025 ). Nonetheless, the synthesis is limited by four key gaps. First, geographic bias persists with 72% of observations originating from southeastern Brazil, leaving important agricultural areas, such as Mato Grosso, Maranhão, Tocantins, Bahia, and other states under‑represented. Second, micronutrients and S were rarely reported, hindering full nutrient‑budget assessments. Third, only 11% of studies tracked biomass decomposition or nutrient release over time, so temporal dynamics remain uncertain. Fourth, there is a notable lack of studies evaluating cover crop mixtures with multiple species and functional diversity. Expansion of research in this area is essential to understand how management strategies of biodiversification can influence not only nutrient and carbon cycling, but also other critical soil functions (e.g., decompaction, biological control of invasive organisms, water retention, thermal regulation, erosion control, and biological nitrogen fixation). Addressing these gaps will require coordinated long‑term trials across diverse edaphoclimatic zones and standardized nutrient‑analysis protocols. That said, this synthesis serves as a baseline for calibrating soil-carbon models, refining nutrient-return targets, and guiding future cover-crop research and policy across Brazil’s varied agroecosystems. Conclusion This systematic review consolidates 1165 field observations and demonstrates that cover crops used in Brazilian agriculture supply substantial biomass (0.33–17.9 Mg ha⁻¹) and deliver appreciable macronutrient recovery (up to 121, 14 and 131 kg ha⁻¹ of N, P and K, respectively). Fabaceae excelled in biological N fixation, Poaceae in biomass production and K scavenging, Brassicaceae in rapid early uptake and soil‑structuring tap‑roots, and Asteraceae in high K returns with prolonged residue persistence. Multispecies cover‑crop mixtures integrate the N‑fixing capacity of legumes, the high‑biomass and K‑scavenging traits of grasses, and the rapid early growth of brassicas, collectively supplying balanced inputs of C, N, P, and K that compensate typical nutrient deficits across diverse tropical and subtropical production systems. By quantifying trait–function linkages across edaphoclimatic zones, the study provides a reproducible data layer for process‑based carbon models, life‑cycle assessments, and site‑specific recommendations. In subtropical Brazil, existing nitrogen fertilization guidelines incorporate the previous crop data, such as type (e.g., grasses, legumes, or fallow) and biomass production, to soil organic matter levels in order to fine-tune recommendations. However, such context-specific approaches remain limited to a few regions. Expanding this data-driven framework nationwide would make fertilization recommendations more precise, tailored to local conditions, and economically efficient. The ranges reported here enable practitioners to calibrate sowing dates, seed ratios, and termination strategies to synchronize nutrient release with cash‑crop demand, reduce reliance on synthetic fertilizers, and enhance SOC sequestration, cornerstones of regenerative agriculture and Brazil’s climate‑smart production goals. Future research should prioritize under‑represented regions, expand micronutrient and S measurements, track residue turnover beyond a single season, and focus on studying multispecies mixtures over single-species cover crops. Long-term, standardized trials are needed to refine these functional benchmarks and advance the adoption of cover crops that enhance resilience and resource efficiency in tropical and subtropical systems. Declarations Funding This work was supported by the São Paulo Research Foundation (FAPESP) as scholarships for Martha Lustosa Carvalho (2022/13531-3) and Lucas Pecci Canisares (2023/08814-9). Bayer and Fundação ABC supported the scholarship of Leonardo de Aro Galera. The National Council for Scientific and Technological Development (CNPq) supported this research via a Research Productivity Fellowship (302249/2025-7) for Maurício Roberto Cherubin. Competing interests The authors have no relevant financial or non-financial interests to disclose. Author Contribution M.L.C., R.S.S., and M.R.C. conceived and designed the study. M.L.C., R.S.S., and M.R.C. developed the methodology. M.L.C., R.S.S., and L.P.C. performed the formal analysis, while M.L.C. and R.S.S. conducted the investigation. Data curation was carried out by M.L.C. and L.P.C. The original draft was prepared by M.L.C., L.P.C., and L.A.G. All authors contributed to the manuscript revision and approved the final version. M.R.C. supervised and administered the project. Acknowledgement We thank the São Paulo Research Foundation (FAPESP) for supporting the work of Martha Lustosa Carvalho (2022/13531-3) and Lucas Pecci Canisares (2023/08814-9), and Bayer and Fundação ABC for supporting the scholarship of Leonardo de Aro Galera. We thank the Center for Carbon Research in Tropical Agriculture/University of São Paulo (CCARBON/USP) (2021/10573-4). Maurício Roberto Cherubin thanks the National Council for Scientific and Technological Development (CNPq) for his Research Productivity Fellowship (302249/2025-7). Data Availability Data is provided within the manuscript and in the supplementary information file. References Ahmad G, Khan A, Khan AA, et al (2021) Biological control: a novel strategy for the control of the plant parasitic nematodes. 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Sci Total Environ 858:159990. https://doi.org/10.1016/j.scitotenv.2022.159990 Vendig I, Guzman A, De La Cerda G, et al (2023) Quantifying direct yield benefits of soil carbon increases from cover cropping. Nat Sustain 6:1125–1134. https://doi.org/10.1038/s41893-023-01131-7 Wu Y, Davis EC, Sohngen BL (2025) Crop rotation and the impact on soil carbon in the U.S. Corn Belt. Carbon Balance Manag 20:6. https://doi.org/10.1186/s13021-025-00293-5 Yang X, Xiong J, Du T, et al (2024) Diversifying crop rotation increases food production, reduces net greenhouse gas emissions and improves soil health. Nat Commun 15:198. https://doi.org/10.1038/s41467-023-44464-9 Yin C, Osborne SL, Lehman RM (2025) Legacy Effects of Cover Crop on Cash Crop Rhizosphere Microbiota in a No-Till Maize–Soybean Cropping System. Phytobiomes J PBIOMES-09-24-0086-R. https://doi.org/10.1094/PBIOMES-09-24-0086-R Yost JL, Schmidt AM, Koelsch R, Schott LR (2022) Effect of swine manure on soil health properties: A systematic review. SSSAJ 86:450–486. https://doi.org/10.1002/saj2.20359 Yousefi M, Dray A, Ghazoul J (2024) Assessing the effectiveness of cover crops on ecosystem services: a review of the benefits, challenges, and trade-offs. Int J Agricult Sustain 22:2335106. https://doi.org/10.1080/14735903.2024.2335106 Additional Declarations No competing interests reported. Supplementary Files Supplementarymaterial.docx Cite Share Download PDF Status: Published Journal Publication published 03 Mar, 2026 Read the published version in Nutrient Cycling in Agroecosystems → Version 1 posted Editorial decision: Revision requested 19 Oct, 2025 Reviews received at journal 23 Sep, 2025 Reviewers agreed at journal 28 Aug, 2025 Reviewers agreed at journal 16 Jul, 2025 Reviews received at journal 13 Jun, 2025 Reviewers agreed at journal 13 Jun, 2025 Reviewers invited by journal 13 Jun, 2025 Editor assigned by journal 04 Jun, 2025 Submission checks completed at journal 02 Jun, 2025 First submitted to journal 30 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6787742","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":471068831,"identity":"00a5bd30-0709-4f86-ad0e-76e323f72bfa","order_by":0,"name":"Martha Lustosa Carvalho","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA60lEQVRIiWNgGAWjYJACZjjrYQODHKoIdsDYDGcmNjAYk64FiAho0W0/+/xxQQ2DPP/sM4YfEnfYpG84f/YAc+Ee3FrMzqQbNs84xmA441yOsUTimbTcDTfyEphnPMOj5UAaYzMPG0MCwxm2BInEtsNALTwGzDwH8Gg5/wyo5R9DgvwZtuQfQC3pBufPENByA2gLbxtDgsEZ5mMgWxIMDuQQ0vKMcTZvn4ThRqAWi8S2NMOZQL8cnoHXYWkMn3m+2cjLnWFsvvGxzUae7/zZg48L8GiBAglkDg8DYQ1ogIdUDaNgFIyCUTDMAQAmU1NtBUaxIQAAAABJRU5ErkJggg==","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":true,"prefix":"","firstName":"Martha","middleName":"Lustosa","lastName":"Carvalho","suffix":""},{"id":471068832,"identity":"5de0c9a3-8800-4a61-9486-a211b81dbce6","order_by":1,"name":"Leonardo Aro Galera","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Leonardo","middleName":"Aro","lastName":"Galera","suffix":""},{"id":471068833,"identity":"2264114a-1a29-4bac-b5be-03a308ea1385","order_by":2,"name":"Raissa Siepman Scholten","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Raissa","middleName":"Siepman","lastName":"Scholten","suffix":""},{"id":471068834,"identity":"d8e18b2a-a9c1-4af4-9adf-28f271ff22c6","order_by":3,"name":"Lucas Pecci Canisares","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Lucas","middleName":"Pecci","lastName":"Canisares","suffix":""},{"id":471068835,"identity":"e5638059-71b2-453b-89f9-d2b181ff861e","order_by":4,"name":"Maurício Roberto Cherubin","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Maurício","middleName":"Roberto","lastName":"Cherubin","suffix":""}],"badges":[],"createdAt":"2025-05-30 22:38:15","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6787742/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6787742/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10705-026-10470-8","type":"published","date":"2026-03-03T15:57:21+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84786576,"identity":"f49fd5c0-81c5-40a0-9775-5535a5af1dc9","added_by":"auto","created_at":"2025-06-17 10:34:57","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":173729,"visible":true,"origin":"","legend":"\u003cp\u003eFlowchart illustrating the search and selection of articles for the systematic review. Numbers in parentheses (n = x) represent the count of articles retrieved at each step. Numbers in parentheses with an apostrophe (n’ = x) indicate the number of observations.\u003c/p\u003e","description":"","filename":"fig1.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/2f4272e89349eecfdf3127a6.jpg"},{"id":84787480,"identity":"9ac58e9a-54e5-4535-b2dd-2471d42ed6b6","added_by":"auto","created_at":"2025-06-17 10:42:57","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1368082,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of the study sites in the context of the Brazilian territory and its main land uses (a) (Souza et al., 2020). Frequency density of the study sites by latitude (b). Percent of study sites by the main Brazilian climates (c) (Alvares et al., 2013). Percent of study sites by soil type (d) (WRB, 2022).\u003c/p\u003e","description":"","filename":"fig2.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/6cde28aa8bdebe2b776197fd.jpg"},{"id":84786577,"identity":"86551b92-fc14-4f01-b28d-4acf662b9c29","added_by":"auto","created_at":"2025-06-17 10:34:57","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":200020,"visible":true,"origin":"","legend":"\u003cp\u003eMean values and number of observations for biomass and nutrient (nitrogen - N, phosphorus - P, and potassium - K) uptake of cover crops from the four main botanical families and main genera in selected studies. Central diagrams indicate values of the two most studied mixtures.\u003c/p\u003e","description":"","filename":"fig3.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/712caff978cbd93b26fab2d6.jpg"},{"id":84786581,"identity":"034c3ea0-2019-412a-8ce0-47fddbc3c275","added_by":"auto","created_at":"2025-06-17 10:34:57","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":177123,"visible":true,"origin":"","legend":"\u003cp\u003eFrequency density representing the concentration of observations at each value, where the area under each curve corresponds to the total number of points in that group of the distribution of cover crop biomass (a), nutrients uptake, N (b), P (c) and K (d), planting month (f), and growing period (g) for \u003cem\u003eFabaceae\u003c/em\u003e, \u003cem\u003ePoaceae\u003c/em\u003e, \u003cem\u003eBrassicaceae\u003c/em\u003eand \u003cem\u003eAsteraceae\u003c/em\u003e cover crop species.\u003c/p\u003e","description":"","filename":"fig4.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/39bb7c426210423856ed5350.jpg"},{"id":84786582,"identity":"361e96cd-22e0-43a1-b5c5-9a8d6fb396f0","added_by":"auto","created_at":"2025-06-17 10:34:57","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":160371,"visible":true,"origin":"","legend":"\u003cp\u003eFrequency density representing the concentration of observations at each value, where the area under each curve corresponds to the total number of points in that group of the distribution of cover crop biomass, nutrients uptake, planting month, and growing period for cover crop species mixtures.\u003c/p\u003e","description":"","filename":"fig5.tif.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/0498b93e229d04cad1c087c4.jpg"},{"id":104250774,"identity":"276bd124-6314-4499-b941-469f989cf25b","added_by":"auto","created_at":"2026-03-09 16:08:15","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2776818,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/37958d2e-1f36-4a31-965f-b3c4afea2e5e.pdf"},{"id":84786579,"identity":"12d95054-7c8a-442c-879a-bba8d2eb91d1","added_by":"auto","created_at":"2025-06-17 10:34:57","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":21185,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarymaterial.docx","url":"https://assets-eu.researchsquare.com/files/rs-6787742/v1/4343d32d473f74ffb3c5fd9e.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Biomass production and nutrient uptake from cover crops in Brazilian agriculture","fulltext":[{"header":"Introduction","content":"\u003cp\u003eConventional agricultural systems simplify natural ecosystems to maximize the provision of one ecosystem service (i.e., provision of food, feed, fiber, and fuel) to the detriment of the provision of other services (Power, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Pereira et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hasan et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Conservation agriculture practices were developed in the last decades as a framework of management strategies to deter, mitigate, and reverse soil organic carbon (SOC) stock depletion (Poeplau and Don, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; He et al., 2025; Wu et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and other ecosystem services in agroecosystems (Lal, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Jian et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). It is based on new and age-old nature-based solutions (NBS) implemented, to various degrees of adoption, in modern agroecosystems, such as organic fertilization, biological control, integrated systems, no-till, and plant diversification (Ahmad et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Yost et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Prairie et al., 2023). Diversification aims to reverse the loss of soil multifunctionality in agroecosystems. It does so by incorporating functional biodiversity through specific groups of plants. This approach helps restore complex biotic interactions. Consequently, these interactions enhance the delivery of multiple ecosystem services in agroecosystems (Daryanto et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yousefi et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePlant diversification can be implemented in agricultural systems by the incorporation of cover crops. Cover crop use is an ancient management practice with multiple potential benefits to soil health, crop yields, and the environment (Koudahe et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Yin et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Cover crops directly or indirectly support the supply of several soil functions and soil-related ecosystem services such as nutrient cycling and provision, water filtering and flow regulation, climate regulation, erosion control, and pest population control (Van Eerd et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Qiu et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Sch\u0026ouml;n et al, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These benefits are related to the capacity of cover crops to increase SOC stocks in agroecosystems (Schlesinger, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In tropical climates, cover cropping can increase SOC by 7.2% on average (Jian et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In the Brazilian context, the edaphoclimatic conditions and management practices from north to south of the country provide a wide range of options for cultivating cover crops throughout the agricultural year (Cherubin et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn tropical agricultural systems, cover crops are often classified into functional groups of families based on the specific ecosystem services they can provide to agroecosystems. For example, tropical grasses can accumulate high biomass in a short period, resulting in biomass left in the field covering the soil surface to prevent erosion and nutrient loss (e.g., Baptistela et al., 2020; Souza et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These species also possess vigorous root systems capable of scavenging water and nutrients at great soil depths and improving subsequent cash crop production (Costa et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, the thick layer of residue can hinder planting operations, and depending on the litter stoichiometry, it can immobilize nutrients, limiting their availability throughout the cash crop growing season. Although legume cover crops typically produce less biomass, they fix atmospheric nitrogen and generate residues with lower C:N ratios, promoting nutrient mineralization and availability during the subsequent cash crop growing season (Vendig et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Peng et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). While cover crops provide numerous beneficial ecosystem services (e.g., Blanco-Canqui et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Daryanto et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yousefi et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), they may also negatively impact other services or crop management practices (e.g., Glaze-Corcoran et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Rouge et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The concurrent growth of cover crops and cash crops in the same space can result in both plants competing for the same resources, even in succession systems (Rouge et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Cover crops are often recommended as a strategy to supply nutrients for the subsequent crop during the litter decomposition; however, the timing of litter decomposition and nutrient release often does not align with the crop\u0026rsquo;s demand (Nevins et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). For instance, grasses with a high C:N ratio can immobilize nitrogen during decomposition, delaying its availability to the following crop and potentially causing nitrogen deficiency and yield reductions (Deines et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Thus, a better understanding of biomass production and nutrient accumulation in the biomass of different groups of cover crops is urgent to provide adequate cover cropping recommendations and avoid or mitigate negative trade-offs. This data can also be used to refine fertilization recommendations, determine the amount of C added to the system, and develop adequate C models or C footprint calculators.\u003c/p\u003e \u003cp\u003eMany challenges regarding cover cropping are still posed to Brazilian researchers, consultants, and farmers. The technical constraints include seed availability, high logistical costs, and access to specialized equipment for cover crop termination. In addition, the scarcity of organized data on biomass production and nutrient extraction by cover crop species across different edaphoclimatic conditions limits the ability of researchers to design more accurate recommendations. Finally, the use of cover crop mixes adds yet another layer of complexity to the questions that remain open for investigation. To fill the basic knowledge gap, in this study, we present an extensive systematic literature review of the available experimental data for cover crops in Brazil, including biomass production and nutrient extraction. The data will assist farmers and consultants in the decision-making process of choosing adequate cover crop species or mixtures for increased soil health in Brazilian farming systems, as well as researchers and policymakers in developing region-specific guidelines, advancing soil management research, and shaping sustainable agricultural practices.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eSearch and dataset preparation\u003c/h2\u003e \u003cp\u003eThe search for articles was restricted by inclusion and exclusion criteria established to find the relevant literature for the systematic review. The inclusion criteria were: i) the publication must be a peer-reviewed article published in a scientific journal; ii) the paper must include cover crop species; iii) the paper must include biomass production, and the uptake by the cover crop of at least one of the following elements: carbon, nitrogen, phosphorus, potassium, calcium, and magnesium; iv) the paper must include the experiment duration with planting and termination dates; v) the studies must be conducted in the field; vi) the experiment must be conducted in Brazil, and vii) the paper must contain the exact or approximate location of the experiment. The exclusion criteria used were: i) studies conducted in semi-perennial systems (e.g., sugarcane), orchards (e.g., oranges, coffee), or vegetable gardens; ii) studies with non-conventional cover crop species (e.g., \u003cem\u003eCalotropis procera\u003c/em\u003e); iii) results reported in non-compatible measurement units (e.g. g/plant, g/pot); and iv) results presented as the average of two or more cropping years.\u003c/p\u003e \u003cp\u003eThe search for articles in the Scopus and Web of Science databases was done using keywords in English. In the Scielo database, the search was in English and Portuguese. The intent of using two languages was to access the highest number of published literature in Brazilian databases, including articles published in academic journals not available in English. The search considered terms mentioned in each record's \"title, abstract, and keywords\" (Topic Field). The search query in English used was: (\"cover crop*\" OR \"green manur*\") AND (\"biomass\" OR \"dry matter\"). The search query in Portuguese for the Scielo database was: (\"planta* de cobertura\" OR \"adub* verde\") AND (\"biomassa\" OR \"mat\u0026eacute;ria seca\"). The search was restricted to studies from Brazil using the \u0026ldquo;country/region\u0026rdquo; filtering tool available on the platforms. The search on the three databases occurred on 21 May 2022. Initially, a total of 1372 papers were retrieved by the search (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All the retrieved articles were organized using reference management software and sorted to remove duplicates. Thus, the number of papers in the dataset was reduced to 840 (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Each unique paper retrieved from the databases was manually sorted by the inclusion/exclusion criteria pre-established before the search. The papers matching all criteria were selected to build the raw dataset used for statistical analysis. The sorting reduced the number of articles from the original 840 to 126 selected articles (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eSubset data within the raw data frame\u003c/h3\u003e\n\u003cp\u003eFrom the 126 articles selected for the raw database, data were extracted on biomass production, scientific names of the cover crop species, geographic coordinates, planting season, experiment duration, soil texture, and soil classification. Data reported in non-standard units were converted to the standard measurement (i.e., kg ha⁻\u0026sup1;). Observations of biomass production below 300 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e or exceeding 18 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e were also removed as they were considered outliers for any cover crop within a field study with a growing period ranging from 30 to 200 days, the latter of which would allow for at least 156 days for one or more cash crop harvest (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The data was organized into a spreadsheet, totaling 1165 individual observations. Using this database, the data were subdivided by grouping each species of cover crop into its respective taxonomic families. After the data filtering, we obtained 758 observations of \u003cem\u003eFabaceae\u003c/em\u003e, \u003cem\u003ePoaceae\u003c/em\u003e, \u003cem\u003eBrassicaceae\u003c/em\u003e, \u003cem\u003eAsteraceae\u003c/em\u003e, and other less common families in monoculture, as well as 199 observations of multispecies mixtures of \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e, two genera or species of \u003cem\u003ePoaceae\u003c/em\u003e, \u003cem\u003eBrassicaceae\u003c/em\u003e, and \u003cem\u003ePoaceae\u003c/em\u003e, and other less common mixtures.\u003c/p\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eThe frequency of observations of biomass, the N, P, and K uptake, planting months, and the growing period were organized in histograms by taxonomic family. An exploratory statistical analysis was performed to obtain each variable's mean, median, and standard deviation using standard statistical libraries from the R software.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eWe sought to obtain an overview of the biomass production, C inputs, and N, P, and K uptake by cover crops from botanical families commonly used in Brazilian agriculture. The main region represented in studies was Central Brazil, with over 72% of total observations and 68% of total unique studies. The next region represented was Southern Brazil, below the Tropic of Capricorn, with 21% of observations and 22% of unique studies (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). In terms of climate, 53% of the study sites are in the humid subtropical regions, 46% in tropical regions, and less than 1% in dry regions (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec). Regarding soil classification, 69% of the sites are on Ferralsols, 17% on Acrisols, 9% on Cambisols, and 5% on other soil types, including Arenosol, Fluvisol, Lixisol, Nitisol, and Plinthosol (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSixty-six species of cover crops and 58 multispecies mixes were retrieved from the 119 articles included in this systematic review. Overall, they belong to seven botanical families, i.e., \u003cem\u003ePoaceae\u003c/em\u003e, \u003cem\u003eFabaceae\u003c/em\u003e, \u003cem\u003eAsteraceae\u003c/em\u003e, \u003cem\u003eBrassicaceae\u003c/em\u003e, \u003cem\u003eAmaranthaceae\u003c/em\u003e, \u003cem\u003ePolygonaceae\u003c/em\u003e, and \u003cem\u003ePedaliaceae\u003c/em\u003e. The \u003cem\u003eAmaranthaceae\u003c/em\u003e, \u003cem\u003ePolygonaceae\u003c/em\u003e, and \u003cem\u003ePedaliaceae\u003c/em\u003e families had fewer than 10 observations each and were grouped as \u0026ldquo;other families\u0026rdquo; for the data analysis. The biomass production ranged from 0.33 to 17.9 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, and the average C content was 44.8% of the total biomass. The C inputs to the system ranged between 489 and 2,898 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, decreasing from \u003cem\u003ePoaceae\u003c/em\u003e (3076 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), \u003cem\u003eAsteraceae\u003c/em\u003e (2781 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), \u003cem\u003eFabaceae\u003c/em\u003e (2393 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), to \u003cem\u003eBrassicaceae\u003c/em\u003e (1727 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) when planted as a single cover crop species. The general N uptake of single cover crop species ranged from 17 to 119 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. The highest mean values were found for the \u003cem\u003eFabaceae\u003c/em\u003e (121 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) family, followed by \u003cem\u003ePoaceae\u003c/em\u003e (107 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), \u003cem\u003eBrassicaceae\u003c/em\u003e (80 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), and \u003cem\u003eAsteraceae\u003c/em\u003e (51 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The mean P uptake of the cover crops ranged from 4.8 to 24 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e from \u003cem\u003ePoaceae\u003c/em\u003e (24.2 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), \u003cem\u003eAsteraceae\u003c/em\u003e (17.5 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), \u003cem\u003eFabaceae\u003c/em\u003e (15.3 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), to \u003cem\u003eBrassicaceae\u003c/em\u003e (4.8 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The K uptake ranged from 33 to 116 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, decreasing from \u003cem\u003ePoaceae\u003c/em\u003e (132 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), \u003cem\u003eAsteraceae\u003c/em\u003e (105 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) and \u003cem\u003eFabaceae\u003c/em\u003e (95 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) to \u003cem\u003eBrassicaceae\u003c/em\u003e (51 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) when planted as single species (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor multispecies mixtures, the C inputs varied from 486 to 2,666 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, decreasing from the \u003cem\u003eFabaceae\u003c/em\u003e with \u003cem\u003ePoaceae\u003c/em\u003e mix (2860 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), to \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e mixes (2461 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), and to the mix of two \u003cem\u003ePoaceae\u003c/em\u003e (1552 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), which resulted in the lowest C inputs. Multispecies mixtures of different families resulted in the highest C inputs among the mixtures. The cover crop mixtures with plants of the \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e (149 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) families, and the \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e mixtures (127 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) had similar N uptake. However, data on \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e mixtures were significantly scarcer than other mixtures (\u003cem\u003en\u0026thinsp;=\u0026thinsp;9\u003c/em\u003e). Data on P and K content were reported for the \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e (17 and 107 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) mix, and the mix of two or more \u003cem\u003ePoaceae\u003c/em\u003e (2 and 31 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The most prevalent cover crop mixtures included plants from the \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e families, accounting for 51% of all observations, followed by mixtures with two or more species of \u003cem\u003ePoaceae\u003c/em\u003e with 31% of observations. To a smaller extent, \u003cem\u003eBrassicaceae\u003c/em\u003e was also represented in studies with cover crop mixtures, with 27 observations of \u003cem\u003eBrassicaceae\u003c/em\u003e species being used in mixtures with plants from the \u003cem\u003ePoaceae\u003c/em\u003e family.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eFabaceae\u003c/h3\u003e\n\u003cp\u003eA total of 82 studies resulted in 415 observations of biomass production of \u003cem\u003eFabaceae\u003c/em\u003e species within the established constraints. A total of 21 genera of plants belonging to the \u003cem\u003eFabaceae\u003c/em\u003e family were found in the dataset and can be consulted in the supplementary material (\u003cb\u003eTable \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003c/b\u003e). Biomass production values for \u003cem\u003eFabaceae\u003c/em\u003e range from 0.33 to 17.85 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with a mean, median, and standard deviation of 5.34, 4.38, and 3.90 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea - blue line). The median extraction of N, P, and K is 105, 9, and 60 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. \u003cem\u003eFabaceae\u003c/em\u003e has a median growth period between planting and desiccation of 104 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef), with planting operations occurring more frequently between either February and June or in November, although the data show planting in all months of the year (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). Species ideally planted in the summer months, like \u003cem\u003eCajanus cajan\u003c/em\u003e (5.9 Mg ha⁻\u0026sup1;) and \u003cem\u003eCrotalaria juncea\u003c/em\u003e (6.7 Mg ha⁻\u0026sup1;) had a median biomass production of 5.0 Mg ha⁻\u0026sup1;, while the winter cover crops, such as \u003cem\u003eLupinus\u003c/em\u003e spp. (4.1 Mg ha⁻\u0026sup1;) and \u003cem\u003eVicia\u003c/em\u003e spp. (2.8 Mg ha⁻\u0026sup1;), had a median of 2.59 Mg ha⁻\u0026sup1; (\u003cb\u003eTable \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003c/b\u003e). In the subtropical region of Brazil, below latitude 23\u0026deg;S, the median biomass production was 5.7 Mg ha⁻\u0026sup1; (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;54) in the summer and 2.88 Mg ha⁻\u0026sup1; (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;15) in the winter.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003ePoaceae\u003c/h2\u003e \u003cp\u003eA total of 84 studies resulted in 422 observations of biomass production for the \u003cem\u003ePoaceae\u003c/em\u003e family. A total of 14 genera of plants belonging to the \u003cem\u003ePoaceae\u003c/em\u003e family were found in the dataset and can be consulted in the supplementary material (\u003cb\u003eTable \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003c/b\u003e). Biomass production of \u003cem\u003ePoaceae\u003c/em\u003e ranged between 0.35 and 17.73 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with a mean, median, and standard deviation of 6.86, 6.21, and 4.13 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively. The median extractions of N, P, and K from \u003cem\u003ePoaceae\u003c/em\u003e are 96, 14, and 99 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e over a mean growth period between planting and desiccation of 109 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef - green line), with most plantings occurring between February and May (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). The biomass production reported for the \u003cem\u003ePoaceae\u003c/em\u003e family spanned from 0.35 to 17.73 Mg ha⁻\u0026sup1;, and a median value of 6.21 Mg ha⁻\u0026sup1;. In the tropical region, the three main grasses planted were \u003cem\u003eCenchrus\u003c/em\u003e spp., with median biomass production of 5.3 Mg ha⁻\u0026sup1;; \u003cem\u003eSorghum bicolor\u003c/em\u003e with 3.8 Mg ha⁻\u0026sup1;; and \u003cem\u003eUrochloa ruziziensis\u003c/em\u003e with 9 Mg ha⁻\u0026sup1;. The most frequently used species below latitude 23\u0026deg;S were \u003cem\u003eAvena\u003c/em\u003e spp. and \u003cem\u003eCenchrus\u003c/em\u003e spp., followed by \u003cem\u003eUrochloa\u003c/em\u003e spp.. The median biomass production of these species was 4 Mg ha⁻\u0026sup1;, 8.6 Mg ha⁻\u0026sup1;, and 1.2 Mg ha⁻\u0026sup1;.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eBrassicaceae\u003c/h3\u003e\n\u003cp\u003eA total of 19 studies resulted in 42 observations of biomass production of \u003cem\u003eBrassicaceae\u003c/em\u003e. We found 2 genera of \u003cem\u003eBrassicaceae\u003c/em\u003e used in the field, namely \u003cem\u003eRaphanus\u003c/em\u003e and \u003cem\u003eCrambe\u003c/em\u003e (\u003cb\u003eTable \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003c/b\u003e). Biomass production of \u003cem\u003eBrassicaceae\u003c/em\u003e ranges from 0.73 to 8.30 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with a mean, median, and standard deviation of 3.85, 3.59, and 1.78 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively. The median extractions of N, P, and K from \u003cem\u003eBrassicaceae\u003c/em\u003e are 66, 6, and 60 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e over a mean growth period between planting and desiccation of 109 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef - purple line), with the most frequent plantings occurring between March and May (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). No studies were observed for planting dates in January, August, September, or December (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). \u003cem\u003eBrassicaceae\u003c/em\u003e exhibited a biomass production ranging from 0.73 to 8.30 Mg ha⁻\u0026sup1;, with a median of 3.59 Mg ha⁻\u0026sup1;. The main species used was \u003cem\u003eRaphanus sativus\u003c/em\u003e, commonly planted as a winter cover crop in the subtropical region. Median P and K uptakes for the family were 6 kg ha⁻\u0026sup1; and 53 kg ha⁻\u0026sup1;. The mean C:N ratio was consistent with the ratio for a localized long-term study with \u003cem\u003eR. sativus\u003c/em\u003e, around 22 (Balota et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eAsteraceae\u003c/h3\u003e\n\u003cp\u003eA total of 6 studies resulted in 14 observations of biomass production for the \u003cem\u003eAsteraceae\u003c/em\u003e family. For the two genera of \u003cem\u003eAsteraceae\u003c/em\u003e used in the field, biomass data were compiled for \u003cem\u003eHelianthus\u003c/em\u003e and \u003cem\u003eThitonia\u003c/em\u003e (\u003cb\u003eTable \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e\u003c/b\u003e). Biomass production of \u003cem\u003eAsteraceae\u003c/em\u003e ranges from 0.90 to 15.20 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with a mean, median, and standard deviation of 6.21, 6.47, and 3.41 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively. The median extractions of N, P, and K from \u003cem\u003eAsteraceae\u003c/em\u003e are 65, 4, and 116 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1,\u003c/sup\u003e respectively, over a mean growth period between planting and desiccation of 112 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ef - red line), with the most frequent plantings occurring between February and March. No studies were observed for plantings in January, April, and between July and December (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ee). The main species used was \u003cem\u003eHelianthus annuus\u003c/em\u003e, also a cash crop planted for vegetable oil production. Median P and K uptakes for the family were 1.7 kg ha⁻\u0026sup1; and 105 kg ha⁻\u0026sup1;. The median C:N ratio for the family was 39.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eCover crop mixtures\u003c/h2\u003e \u003cp\u003eThe most used mixtures in the selected studies were combinations of \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e species, with 32 studies resulting in 256 observations. The biomass production of the mixtures of \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e species ranged between 0.63 and 17.00 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with the mean, median, and standard deviation of 6.38, 5.71, and 3.45 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea - blue line). The median of the N, P, and K extractions were 104, 11, and 80 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1,\u003c/sup\u003e with an average planting period between planting and termination of 107 days, and the most frequent planting months were between March, May, and November (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ee,f). The most common mixture for the subtropical region of Brazil was \u003cem\u003eAvena strigosa\u003c/em\u003e with \u003cem\u003eVicia sativa\u003c/em\u003e, and \u003cem\u003eCajanus cajan\u003c/em\u003e with \u003cem\u003eUrochloa ruziziensis\u003c/em\u003e for the tropical region. Medians of biomass production for the subtropical and tropical regions were 4.4 Mg ha⁻\u0026sup1; and 6.6 Mg ha⁻\u0026sup1;. We found similar P uptakes for both regions, with 14.2 Mg ha⁻\u0026sup1; and 13.6 Mg ha⁻\u0026sup1; for the subtropical and tropical regions. The median K uptakes were also similar, 121 Mg ha⁻\u0026sup1; and 103 Mg ha⁻\u0026sup1; for the subtropical and tropical regions. The C:N ratio for the subtropical region with \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e species was 22.8, while tropical mixtures had a higher C:N ratio of 40.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSeven studies with a total of 27 observations presented data for mixtures of \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e species. We identified 5 different combinations of species of the two families, all of them with \u003cem\u003eRaphanus sativus\u003c/em\u003e. The biomass production of the mixtures of \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e ranged between 0.64 and 10.40 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with the mean, median, and standard deviation of 5.49, 5.22, and 2.74 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively. The median of the N extraction was 106 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with the average growing period between planting and termination of 107 days, and the most frequent planting months were between March and May (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ee,f - purple line). No P and K concentrations were reported for mixtures of \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e in the studies retrieved and selected in this systematic review.\u003c/p\u003e \u003cp\u003eMixtures of two \u003cem\u003ePoaceae\u003c/em\u003e species were identified in 8 studies with 67 total observations. We identified 11 different combinations of species. The biomass production of these mixtures ranged between 0.33 and 16.40 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with the mean, median, and standard deviation of 3.46, 1.09 e 4.10 Mg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively. The medians of the N, P, and K extraction were 17, 1, and 33 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e with the growing period between planting and termination of 120 days, and the planting months were mostly between March and April (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ee,f - green line).\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eBiomass production and nutrients uptake\u003c/h2\u003e \u003cp\u003eThis systematic literature review provides insights into biomass production and nutrient extraction of cover crops in Brazilian agroecosystems, linking family traits to ecosystem services. \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e families dominated the studies, accounting for over 92% of the observations. Species from the genera \u003cem\u003eUrochloa\u003c/em\u003e, \u003cem\u003eCrotalaria\u003c/em\u003e, and \u003cem\u003eCenchrus\u003c/em\u003e represented 40% of these occurrences, largely due to their proven effectiveness in improving soil health and nutrient cycling, as well as their hardiness and strong adaptation to Brazilian climatic and soil conditions.\u003c/p\u003e \u003cp\u003eAs the highest biomass-producing family of cover crops, species of \u003cem\u003ePoaceae\u003c/em\u003e achieved a median production of 6.21 Mg ha⁻\u0026sup1;. This result supports earlier reports that tropical grasses excel in biomass production thanks to efficient nutrient uptake and deep, extensive roots, traits well documented for \u003cem\u003eUrochloa ruziziensis\u003c/em\u003e (Baptistella et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). \u003cem\u003eFabaceae\u003c/em\u003e displayed moderately high biomass (median 4.38 Mg ha⁻\u0026sup1;), underscoring their dual contribution of substantial biomass alongside N fixation capabilities, particularly notable in tropical legumes such as \u003cem\u003eCrotalaria juncea\u003c/em\u003e and \u003cem\u003eCajanus cajan\u003c/em\u003e (Berriel and Perdomo, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The intermediate productivity observed in \u003cem\u003eBrassicaceae\u003c/em\u003e (median 3.59 Mg ha⁻\u0026sup1;) aligns with their recognized rapid growth but limited overall biomass compared to grasses (Elhakeem et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Interestingly, \u003cem\u003eAsteraceae\u003c/em\u003e also presented high biomass yields (median 6.47 Mg ha⁻\u0026sup1;), higher than most species of the \u003cem\u003ePoaceae\u003c/em\u003e family and indicating variability possibly influenced by species selection, climate conditions, and management strategies across studies (Freitas et al., 2019).\u003c/p\u003e \u003cp\u003eTropical grasses like \u003cem\u003eUrochloa spp.\u003c/em\u003e are preferred in the Brazilian savannah for rapid growth, drought tolerance, and high biomass yield, whereas in subtropical areas, grasses like ryegrass dominate due to cold tolerance and forage potential (Pavinato et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Baptistella et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, biomass yields of tropical grasses can be lower in subtropical conditions, as shown by median biomass production of \u003cem\u003eU. brizantha\u003c/em\u003e (6.5 Mg ha\u0026sup1; in tropical; 3.4 Mg ha\u0026sup1; in subtropical), emphasizing the necessity for selecting species suitable for local edaphoclimatic conditions to optimize agroecosystem benefits.\u003c/p\u003e \u003cp\u003eFurthermore, cover crops classified as \u003cem\u003eFabaceae\u003c/em\u003e, particularly species of \u003cem\u003eCrotalaria\u003c/em\u003e, are strategically used by farmers to control plant-parasitic nematodes sustainably, an issue increasingly critical under climate change conditions, which may enhance nematode survival rates (Lopes et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Khanal and Land, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Souza et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Moreover, \u003cem\u003eFabaceae\u003c/em\u003e exhibited the highest N content and lowest C:N ratio (median C:N ratio of 16.9), consistent with studies in Brazilian savannah agroecosystems (Carvalho et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough individual species can provide ecosystem services, the combination of more than one species can balance different services, such as mixing grasses with legumes for balancing the high biomass production of the first and the residue quality (lower C:N) of the second (Plumhoff et al., 2022). In this study, the use of multispecies mixtures followed similar single-species trends and provided complementary functions and broadened ecosystem services. \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e combinations made up 57% of observations, followed by mixtures with multiple \u003cem\u003ePoaceae\u003c/em\u003e species and \u003cem\u003ePoaceae\u003c/em\u003e cover crops intercropped with cash crops.\u003c/p\u003e \u003cp\u003eThe most common use of mixtures of \u003cem\u003ePoaceae\u003c/em\u003e and \u003cem\u003eFabaceae\u003c/em\u003e species can be explained by plant complementary traits and functions (Schipanski et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The mixtures incorporate both the N-fixing capabilities of \u003cem\u003eFabaceae\u003c/em\u003e species with the high biomass potential of \u003cem\u003ePoaceae\u003c/em\u003e. The goal of using these mixtures then is to provide greater soil cover with lower N immobilization for the subsequent cash crop, and higher C accrual. This approach also aligns with the understanding that effective soil C sequestration depends on balancing C inputs with N availability, as soil organic matter formation and stabilization require proportionally more N than plant biomass (Cotrufo et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Moreover, mixtures of \u003cem\u003ePoaceae\u003c/em\u003e and \u003cem\u003eFabaceae\u003c/em\u003e have been shown to improve soil health and reduce yield variability over time, promoting greater resilience to climate-related stresses (Souza et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Yet the use of mixtures demands greater management expertise from farmers. Per our results, biomass production for \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e mixtures varied from 0.63 Mg ha⁻\u0026sup1; to 17 Mg ha⁻\u0026sup1;, with an overall median of 5.7 Mg ha⁻\u0026sup1;. The range is due to differences in climatic conditions, planting seasons, and species of choice, the same as for single species. In addition, initial seeding proportions of mixtures, when available in the consulted literature, were highly variable and could have influenced the evaluated variables. Our results also revealed that mixtures used in the subtropical region, usually planted in winter, allow for the preponderance of \u003cem\u003eFabaceae\u003c/em\u003e plants with lower C:N ratios, while mixtures used in the tropics have C:N ratios much more similar to general \u003cem\u003ePoaceae\u003c/em\u003e, indicating a preponderance of these plants in the mixture.\u003c/p\u003e \u003cp\u003eIn Brazil, intercropping cover crops with cash crops (Santa F\u0026eacute; System) using \u003cem\u003eUrochloa\u003c/em\u003e grasses (mainly \u003cem\u003eU. ruziziensis\u003c/em\u003e and \u003cem\u003eU. brizantha\u003c/em\u003e) is common, primarily to protect soil from erosive tropical rainfall events. The mean biomass produced by \u003cem\u003eUrochloa\u003c/em\u003e grasses is approximately 5.7 Mg ha⁻\u0026sup1;, varying according to the cash crop due to competition for light and nutrients. Biomass production is higher with soybean (5.3 Mg ha⁻\u0026sup1;) than with maize (4.19 Mg ha⁻\u0026sup1;), while intercropping with common beans achieves 7.9 Mg ha⁻\u0026sup1;, although studies with beans are limited. While intercropping usually does not significantly impact maize yield (Brito et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Souza et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), soybean yields can be negatively affected if not properly managed, necessitating selective herbicide use (Saraiva et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Additionally, intercropped \u003cem\u003eUrochloa\u003c/em\u003e grasses facilitate integrated crop-livestock systems by providing forage during dry winters, enhancing livestock productivity, and diversifying farmers' income (Souza et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBeyond biomass production, the same families differed markedly in their ability to capture and recycle key nutrients. \u003cem\u003eFabaceae\u003c/em\u003e fix abundant N (median 105 kg ha⁻\u0026sup1;) yet yield moderate biomass, whereas \u003cem\u003ePoaceae\u003c/em\u003e generate the greatest biomass and recover similar N (96 kg ha⁻\u0026sup1;) but much more K (99 kg ha⁻\u0026sup1;), a reflection of deep roots and rapid growth (Passot et al., 2016). \u003cem\u003eBrassicaceae\u003c/em\u003e show swift early uptake, removing 66 kg N and 60 kg K ha⁻\u0026sup1; within about 109 days while alleviating compaction via tap-roots (Tribouillois et al., 2015). Their rapid growth makes them excellent plants to cover the short fallow periods between harvests in the Brazilian crop succession systems common in the subtropical South and irrigated areas of western Bahia (Altieri et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). \u003cem\u003eAsteraceae\u003c/em\u003e, dominated by sunflower in this review, accumulate high K (116 kg ha⁻\u0026sup1;) but lower N and P, and the high C:N ratio (\u0026gt;\u0026thinsp;40) suggests slower mineralization and longer soil cover. Mixtures such as millet and crotalaria synergize N fixation with K capture, boosting total nutrient return and subsequent crop yields over monocultures (Silva et al., 2010; Souza et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Phosphorus and K uptake by cover crops significantly varied between subtropical and tropical regions, reflecting regional differences in species selection and biomass accumulation.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eImplications\u003c/h2\u003e \u003cp\u003eWhen biomass and nutrient profiles are viewed together, clear agronomic patterns emerge that inform agronomic decisions for optimizing agricultural practices and sustainable soil management strategies. Biomass production varies widely (0.33 to 17.9 Mg ha⁻\u0026sup1;), with average C inputs by families ranging from 489 to 2,898 kg ha⁻\u0026sup1;. This information is especially relevant for modeling soil carbon stocks and estimating the carbon footprint of agricultural systems (Jordon et al., 2022; McClelland et al., 2021; Prairie et al., 2023). Additionally, cover crops play a critical role in climate adaptation by leaving a protective mulch layer on the soil surface, especially when species with high C:N ratios are used. This mulch moderates diurnal temperature fluctuations and reduces evaporation, creating a more stable microclimate that enhances the resilience of subsequent cash crops to heat and drought stress (Souza et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In particular, \u003cem\u003ePoaceae\u003c/em\u003e species such as \u003cem\u003eUrochloa ruziziensis\u003c/em\u003e and \u003cem\u003eSorghum bicolor\u003c/em\u003e are known for their high biomass production (median of 6.21 Mg ha⁻\u0026sup1;) and high C:N ratios, which contribute to slower decomposition and prolonged soil cover (Souza et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Carvalho et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). With carbon inputs ranging from 489 to 2,898 kg C ha⁻\u0026sup1;, many cover crop species, especially in mixtures, provide enough biomass to match or substantially reduce the typical C deficits observed in degraded tropical soils which can vary from 100 kg C ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e y\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e to 3150 kg C ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e y\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (Medeiros et al., 2021; Oliveira et al., 2022). These inputs are particularly important because they help replenish particulate organic carbon (POC), the soil carbon fraction most vulnerable to degradation under intensive agriculture. As POC plays a key role in soil structure, nutrient retention, and microbial activity, its restoration through high-biomass regenerative practices is essential for rebuilding soil health and long-term productivity (Prairie et al., 2023). Furthermore, biological nitrogen fixation by \u003cem\u003eFabaceae\u003c/em\u003e can reduce the need for external N inputs (Schipanski et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Skinner et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This reduction is particularly relevant for climate mitigation, as the production and application of synthetic nitrogen fertilizers are major sources of greenhouse gas emissions, especially nitrous oxide (N₂O), which has a global warming potential 265 times greater than CO₂ (IPCC, 2021).\u003c/p\u003e \u003cp\u003eThese results also provide valuable insights for designing multi-species cover crop mixtures in Brazil. By quantifying family-specific nutrient uptake and identifying a median growth window of 104 to 112 days, the findings support mixtures that combine N-fixing \u003cem\u003eFabaceae\u003c/em\u003e with high-biomass \u003cem\u003ePoaceae\u003c/em\u003e, supplemented by smaller proportions of \u003cem\u003eBrassicaceae\u003c/em\u003e for pest suppression. These combinations can be timed to regional climatic windows so that sowing dates maximize biomass and ensure that residue decomposition and N release coincide with cash‑crop demand. The data highlights the strong performance of \u003cem\u003ePoaceae\u003c/em\u003e in nutrient uptake, particularly phosphorus (24 kg ha⁻\u0026sup1;) and potassium (131 kg ha⁻\u0026sup1;), underscoring their key role in enhancing nutrient cycling. When combined with other functional groups, such as \u003cem\u003eFabaceae\u003c/em\u003e and \u003cem\u003eBrassicaceae\u003c/em\u003e, these grasses contribute to more balanced cover crop mixtures that support efficient nutrient use and reduce the need for supplementary chemical fertilization (Souza et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eNonetheless, the synthesis is limited by four key gaps. First, geographic bias persists with 72% of observations originating from southeastern Brazil, leaving important agricultural areas, such as Mato Grosso, Maranh\u0026atilde;o, Tocantins, Bahia, and other states under‑represented. Second, micronutrients and S were rarely reported, hindering full nutrient‑budget assessments. Third, only 11% of studies tracked biomass decomposition or nutrient release over time, so temporal dynamics remain uncertain. Fourth, there is a notable lack of studies evaluating cover crop mixtures with multiple species and functional diversity. Expansion of research in this area is essential to understand how management strategies of biodiversification can influence not only nutrient and carbon cycling, but also other critical soil functions (e.g., decompaction, biological control of invasive organisms, water retention, thermal regulation, erosion control, and biological nitrogen fixation). Addressing these gaps will require coordinated long‑term trials across diverse edaphoclimatic zones and standardized nutrient‑analysis protocols. That said, this synthesis serves as a baseline for calibrating soil-carbon models, refining nutrient-return targets, and guiding future cover-crop research and policy across Brazil\u0026rsquo;s varied agroecosystems.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis systematic review consolidates 1165 field observations and demonstrates that cover crops used in Brazilian agriculture supply substantial biomass (0.33\u0026ndash;17.9 Mg ha⁻\u0026sup1;) and deliver appreciable macronutrient recovery (up to 121, 14 and 131 kg ha⁻\u0026sup1; of N, P and K, respectively). \u003cem\u003eFabaceae\u003c/em\u003e excelled in biological N fixation, \u003cem\u003ePoaceae\u003c/em\u003e in biomass production and K scavenging, \u003cem\u003eBrassicaceae\u003c/em\u003e in rapid early uptake and soil‑structuring tap‑roots, and \u003cem\u003eAsteraceae\u003c/em\u003e in high K returns with prolonged residue persistence. Multispecies cover‑crop mixtures integrate the N‑fixing capacity of legumes, the high‑biomass and K‑scavenging traits of grasses, and the rapid early growth of brassicas, collectively supplying balanced inputs of C, N, P, and K that compensate typical nutrient deficits across diverse tropical and subtropical production systems. By quantifying trait\u0026ndash;function linkages across edaphoclimatic zones, the study provides a reproducible data layer for process‑based carbon models, life‑cycle assessments, and site‑specific recommendations. In subtropical Brazil, existing nitrogen fertilization guidelines incorporate the previous crop data, such as type (e.g., grasses, legumes, or fallow) and biomass production, to soil organic matter levels in order to fine-tune recommendations. However, such context-specific approaches remain limited to a few regions. Expanding this data-driven framework nationwide would make fertilization recommendations more precise, tailored to local conditions, and economically efficient.\u003c/p\u003e \u003cp\u003eThe ranges reported here enable practitioners to calibrate sowing dates, seed ratios, and termination strategies to synchronize nutrient release with cash‑crop demand, reduce reliance on synthetic fertilizers, and enhance SOC sequestration, cornerstones of regenerative agriculture and Brazil\u0026rsquo;s climate‑smart production goals. Future research should prioritize under‑represented regions, expand micronutrient and S measurements, track residue turnover beyond a single season, and focus on studying multispecies mixtures over single-species cover crops. Long-term, standardized trials are needed to refine these functional benchmarks and advance the adoption of cover crops that enhance resilience and resource efficiency in tropical and subtropical systems.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work was supported by the S\u0026atilde;o Paulo Research Foundation (FAPESP) as scholarships for Martha Lustosa Carvalho (2022/13531-3) and Lucas Pecci Canisares (2023/08814-9). Bayer and Funda\u0026ccedil;\u0026atilde;o ABC supported the scholarship of Leonardo de Aro Galera. The National Council for Scientific and Technological Development (CNPq) supported this research via a Research Productivity Fellowship (302249/2025-7) for Maur\u0026iacute;cio Roberto Cherubin.\u003c/p\u003e \u003cp\u003eCompeting interests\u003c/p\u003e \u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eM.L.C., R.S.S., and M.R.C. conceived and designed the study. M.L.C., R.S.S., and M.R.C. developed the methodology. M.L.C., R.S.S., and L.P.C. performed the formal analysis, while M.L.C. and R.S.S. conducted the investigation. Data curation was carried out by M.L.C. and L.P.C. The original draft was prepared by M.L.C., L.P.C., and L.A.G. All authors contributed to the manuscript revision and approved the final version. M.R.C. supervised and administered the project.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe thank the S\u0026atilde;o Paulo Research Foundation (FAPESP) for supporting the work of Martha Lustosa Carvalho (2022/13531-3) and Lucas Pecci Canisares (2023/08814-9), and Bayer and Funda\u0026ccedil;\u0026atilde;o ABC for supporting the scholarship of Leonardo de Aro Galera. We thank the Center for Carbon Research in Tropical Agriculture/University of S\u0026atilde;o Paulo (CCARBON/USP) (2021/10573-4). Maur\u0026iacute;cio Roberto Cherubin thanks the National Council for Scientific and Technological Development (CNPq) for his Research Productivity Fellowship (302249/2025-7).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData is provided within the manuscript and in the supplementary information file.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAhmad G, Khan A, Khan AA, et al (2021) Biological control: a novel strategy for the control of the plant parasitic nematodes. Antonie van Leeuwenhoek 114:885\u0026ndash;912. https://doi.org/10.1007/s10482-021-01577-9\u003c/li\u003e\n\u003cli\u003eAltieri MA, Lana MA, Bittencourt HV, et al (2011) Enhancing Crop Productivity via Weed Suppression in Organic No-Till Cropping Systems in Santa Catarina, Brazil. 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Int J Agricult Sustain 22:2335106. https://doi.org/10.1080/14735903.2024.2335106\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"nutrient-cycling-in-agroecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"fres","sideBox":"Learn more about [Nutrient Cycling in Agroecosystems](http://link.springer.com/journal/10705)","snPcode":"10705","submissionUrl":"https://submission.nature.com/new-submission/10705/3","title":"Nutrient Cycling in Agroecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"green manure, soil health, regenerative agriculture, functional biodiversity, nutrient cycling","lastPublishedDoi":"10.21203/rs.3.rs-6787742/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6787742/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eIn a scenario of global soil degradation from inadequate management, cover crops are increasingly promoted for their positive effects on soil health and ecosystem services. In Brazil, however, their use remains limited due to technical barriers and a lack of accessible data. In this systematic literature review, we extensively searched for data on biomass production and nutrient uptake of the primary cover crop families used in Brazilian agriculture. We aimed to map available research, identify knowledge gaps, and offer data to inform cover crop adoption. We retrieved over 1,300 articles from three databases, selecting 126 studies that yielded more than 1,000 observations of cover crops grown as single species or in mixtures. In total, 66 species and 58 multispecies mixes from seven botanical families were represented. Cover crop biomass production ranged from 0.33 to 17.9 Mg ha⁻\u0026sup1;, with an average carbon content of 44.8%, and C inputs varying from 489 to 2,898 kg ha⁻\u0026sup1;, decreasing from \u003cem\u003ePoaceae\u003c/em\u003e to \u003cem\u003eBrassicaceae\u003c/em\u003e. Nitrogen uptake ranged from 17 to 119 kg ha⁻\u0026sup1;, with the highest values in \u003cem\u003eFabaceae\u003c/em\u003e and similar uptake in \u003cem\u003eBrassicaceae\u003c/em\u003e and \u003cem\u003ePoaceae\u003c/em\u003e mixtures. Phosphorus and potassium uptake varied across species, with \u003cem\u003ePoaceae\u003c/em\u003e showing the highest phosphorus and potassium, while mixtures had varied nutrient content, with limited data on phosphorus and potassium. Overall, these findings highlight the variability in nutrient uptake among different cover crop species and mixtures, providing valuable insights for optimizing cover crop selection to enhance soil health and nutrient management.\u003c/p\u003e","manuscriptTitle":"Biomass production and nutrient uptake from cover crops in Brazilian agriculture","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-17 10:34:53","doi":"10.21203/rs.3.rs-6787742/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-10-19T08:18:22+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-09-23T17:48:40+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"241852830648148850206776746638638520724","date":"2025-08-28T12:48:39+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"15864786134517321075027988677859222931","date":"2025-07-16T14:50:39+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-13T19:51:23+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"279850829114042495644614065210262395565","date":"2025-06-13T12:50:12+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-13T07:36:09+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-04T08:56:08+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-06-03T02:28:46+00:00","index":"","fulltext":""},{"type":"submitted","content":"Nutrient Cycling in Agroecosystems","date":"2025-05-30T22:33:16+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"nutrient-cycling-in-agroecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"fres","sideBox":"Learn more about [Nutrient Cycling in Agroecosystems](http://link.springer.com/journal/10705)","snPcode":"10705","submissionUrl":"https://submission.nature.com/new-submission/10705/3","title":"Nutrient Cycling in Agroecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"cfed7d1a-d12c-4a65-9dce-387dc8b756e8","owner":[],"postedDate":"June 17th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-03-09T16:04:29+00:00","versionOfRecord":{"articleIdentity":"rs-6787742","link":"https://doi.org/10.1007/s10705-026-10470-8","journal":{"identity":"nutrient-cycling-in-agroecosystems","isVorOnly":false,"title":"Nutrient Cycling in Agroecosystems"},"publishedOn":"2026-03-03 15:57:21","publishedOnDateReadable":"March 3rd, 2026"},"versionCreatedAt":"2025-06-17 10:34:53","video":"","vorDoi":"10.1007/s10705-026-10470-8","vorDoiUrl":"https://doi.org/10.1007/s10705-026-10470-8","workflowStages":[]},"version":"v1","identity":"rs-6787742","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6787742","identity":"rs-6787742","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}