Nadia I. Maaroufi
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Previous Organizations / Institutes
The organizations / institutes from the scholar's other articles in the past ten years
Research Subjects
Nadia I. Maaroufi has a total of 98 co-authors. Here are the top co-authors by number of shared publications:
Annika Nordin
Department of Forest Genetics and Plant Physiology, Umeå Plant Science Center, Swedish University of Agricultural Sciences, Umeå, Sweden
6 shared publicationsKristin Palmqvist
Department of Ecology and Environmental Science, Umeå University, Umeå, Sweden
4 shared publicationsMichael J. Gundale
Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, Umeå, Sweden
4 shared publications- Michael J. Gundale
Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, Umeå, Sweden
3 shared publications - Benjamin Forsmark
Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden
2 shared publications - Jonathan Richard De Long
Department of Terrestrial Ecology, Netherlands Institute of Ecology, Wageningen, The Netherlands
2 shared publications
Latest Publications
- Open Access
- 18 May 2026
- Published by Elsevier in Agriculture, Ecosystems & Environment
- Vol. 409
Native non-N₂-fixing and exotic N₂-fixing tree species provide higher-quality green manure than exotic fast-growing species in sub-Saharan Africa
- 18 May 2026
- Published by Elsevier in Agriculture, Ecosystems & Environment
- Vol. 409
Open AccessNative non-N₂-fixing and exotic N₂-fixing tree species provide higher-quality green manure than exotic fast-growing species in sub-Saharan Africa
- Open Access
- 16 May 2026
- Published by Elsevier in Data in Brief
- Vol. 66
SLU long-term field experiments: Soil organic matter in a cereal only cropping system (R3-0020), crop and soil data from 1971 to 2023
- 16 May 2026
- Published by Elsevier in Data in Brief
- Vol. 66
Open AccessSLU long-term field experiments: Soil organic matter in a cereal only cropping system (R3-0020), crop and soil data from 1971 to 2023
- 16 March 2026
- Published by Taylor & Francis
- p. 47-62
Soil symphony
Here, we discuss the current and emerging themes in soil processes related to grassland nutrient cycling. Soil processes involve four major components, namely addition, loss, translocation, and transformation of nutrients. The ability of grasslands to provide ecosystem services such as fodder for livestock and carbon storage is intimately linked to how nutrients cycle through the soil and between the greater global stocks of nutrients. We discuss macro- and micronutrients and conclude by identifying future avenues of research that require attention.
- 16 March 2026
- Published by Taylor & Francis
- p. 47-62
Soil symphony
Here, we discuss the current and emerging themes in soil processes related to grassland nutrient cycling. Soil processes involve four major components, namely addition, loss, translocation, and transformation of nutrients. The ability of grasslands to provide ecosystem services such as fodder for livestock and carbon storage is intimately linked to how nutrients cycle through the soil and between the greater global stocks of nutrients. We discuss macro- and micronutrients and conclude by identifying future avenues of research that require attention.
- Open Access
- 19 February 2026
- Published by Springer Nature in Scientific Data
- Vol. 13 (1)
Data from long-term experiments in temperate croplands to evaluate soil organic carbon models
Soil organic carbon (SOC) models need independent evaluation against field measurements, but those latter are rarely publicly available and harmonized. In this study, we collected and shared data from 167 agronomic treatments in 34 agronomic long-term experiments (LTEs) located in temperate croplands, allowing the evaluation of several soil organic C models such as RothC, Century, AMG, MIMICS, ICBM, Millenial, and CTOOL. The dataset includes climate data, soil properties, C inputs from crops (n = 4588 records) and organic amendments, irrigation data, monthly soil cover, as well as SOC stock measurements in the topsoil layer (n = 1328 records). Climate, soil moisture, and soil temperature data were extracted from daily climate databases. Carbon inputs from crops were calculated from observed yields and harvest index, with some harvest index values estimated, combined with crop allometric coefficients from the literature. Descriptions of LTE, agronomic treatments, methodological metadata, and a part of the code, accompanies the dataset. The dataset can be reused to evaluate single SOC models, or to evaluate an ensemble of models.
- 19 February 2026
- Published by Springer Nature in Scientific Data
- Vol. 13 (1)
Open AccessData from long-term experiments in temperate croplands to evaluate soil organic carbon models
Soil organic carbon (SOC) models need independent evaluation against field measurements, but those latter are rarely publicly available and harmonized. In this study, we collected and shared data from 167 agronomic treatments in 34 agronomic long-term experiments (LTEs) located in temperate croplands, allowing the evaluation of several soil organic C models such as RothC, Century, AMG, MIMICS, ICBM, Millenial, and CTOOL. The dataset includes climate data, soil properties, C inputs from crops (n = 4588 records) and organic amendments, irrigation data, monthly soil cover, as well as SOC stock measurements in the topsoil layer (n = 1328 records). Climate, soil moisture, and soil temperature data were extracted from daily climate databases. Carbon inputs from crops were calculated from observed yields and harvest index, with some harvest index values estimated, combined with crop allometric coefficients from the literature. Descriptions of LTE, agronomic treatments, methodological metadata, and a part of the code, accompanies the dataset. The dataset can be reused to evaluate single SOC models, or to evaluate an ensemble of models.
- 22 July 2023
- Published by Wiley in New Phytologist
- Vol. 240 (1) , 399-411
Host plant height explains the effect of nitrogen enrichment on arbuscular mycorrhizal fungal communities
Summary: Nitrogen (N) enrichment is widely known to affect the root‐associated arbuscular mycorrhizal fungal (AMF) community in different ways, for example, via altering soil properties and/or shifting host plant functional structure. However, empirical knowledge of their relative importance is still lacking. Using a long‐term N addition experiment, we measured the AMF community taxonomic and phylogenetic diversity at the single plant species (roots of 15 plant species) and plant community (mixed roots) levels. We also measured four functional traits of 35 common plant species along the N addition gradient. We found divergent responses of AMF diversity to N addition for host plants with different innate heights (i.e. plant natural height under unfertilized treatment). Furthermore, our data showed that species‐specific responses of AMF diversity to N addition were negatively related to the change in maximum plant height. When scaling up to the community level, N addition affected AMF diversity mainly through increasing the maximum plant height, rather than altering soil properties. Our results highlight the importance of plant height in driving AMF community dynamics under N enrichment at both species and community levels, thus providing important implications for understanding the response of AMF diversity to anthropogenic N deposition.
- 22 July 2023
- Published by Wiley in New Phytologist
- Vol. 240 (1) , 399-411
Host plant height explains the effect of nitrogen enrichment on arbuscular mycorrhizal fungal communities
Summary: Nitrogen (N) enrichment is widely known to affect the root‐associated arbuscular mycorrhizal fungal (AMF) community in different ways, for example, via altering soil properties and/or shifting host plant functional structure. However, empirical knowledge of their relative importance is still lacking. Using a long‐term N addition experiment, we measured the AMF community taxonomic and phylogenetic diversity at the single plant species (roots of 15 plant species) and plant community (mixed roots) levels. We also measured four functional traits of 35 common plant species along the N addition gradient. We found divergent responses of AMF diversity to N addition for host plants with different innate heights (i.e. plant natural height under unfertilized treatment). Furthermore, our data showed that species‐specific responses of AMF diversity to N addition were negatively related to the change in maximum plant height. When scaling up to the community level, N addition affected AMF diversity mainly through increasing the maximum plant height, rather than altering soil properties. Our results highlight the importance of plant height in driving AMF community dynamics under N enrichment at both species and community levels, thus providing important implications for understanding the response of AMF diversity to anthropogenic N deposition.
- Open Access
- 9 February 2023
- Published by Wiley in Functional Ecology
- Vol. 37 (3) , 564-575
Plant attributes interact with fungal pathogens and nitrogen addition to drive soil enzymatic activities and their temporal variation
Nitrogen enrichment can alter soil communities and their functioning directly, via changes in nutrient availability and stoichiometry, or indirectly, by changing plant communities or the abundance of consumers. However, most studies have only focused on one of these potential drivers and we know little about the relative importance of the different mechanisms (changes in nutrient availability, in plant diversity or functional composition or in consumer abundance) by which nitrogen enrichment affects soil functioning. In addition, soil functions could vary dramatically between seasons; however, they are typically measured only once during the peak growing season. We therefore know little about the drivers of intra‐annual stability in soil functioning. In this study, we measured activities of β‐glucosidase and acid phosphatase, two extracellular enzymes that indicate soil functioning. We did so in a large grassland experiment which tested the effects, and relative importance, of nitrogen enrichment, plant functional composition and diversity, and foliar pathogen presence (controlled by fungicide) on soil functioning. We measured the activity of the two enzymes across seasons and years to assess the stability and temporal dynamics of soil functioning. Overall β‐glucosidase activity was slightly increased by nitrogen enrichment over time but did not respond to the other experimental treatments. Conversely, plant functional diversity and interactions between plant attributes and fungicide application were important drivers of mean acid phosphatase activity. The temporal stability of both soil enzymes was differently affected by two facets of plant diversity: species richness increased temporal stability and functional diversity decreased it; however, these effects were dampened when nitrogen and fungicide were added. The fungicide effects on soil enzyme activities suggest that foliar pathogens can also affect below‐ground processes and the interacting effect of fungicide and plant diversity suggests that these plant enemies can modulate the relationship between plant diversity and ecosystem functioning. The contrasting effects of our treatments on the mean versus stability of soil enzyme activities clearly show the need to consider temporal dynamics in below‐ground processes, to better understand the responses of soil microbes to environmental changes such as nutrient enrichment. Read the free Plain Language Summary for this article on the Journal blog.
- 9 February 2023
- Published by Wiley in Functional Ecology
- Vol. 37 (3) , 564-575
Open AccessPlant attributes interact with fungal pathogens and nitrogen addition to drive soil enzymatic activities and their temporal variation
Nitrogen enrichment can alter soil communities and their functioning directly, via changes in nutrient availability and stoichiometry, or indirectly, by changing plant communities or the abundance of consumers. However, most studies have only focused on one of these potential drivers and we know little about the relative importance of the different mechanisms (changes in nutrient availability, in plant diversity or functional composition or in consumer abundance) by which nitrogen enrichment affects soil functioning. In addition, soil functions could vary dramatically between seasons; however, they are typically measured only once during the peak growing season. We therefore know little about the drivers of intra‐annual stability in soil functioning. In this study, we measured activities of β‐glucosidase and acid phosphatase, two extracellular enzymes that indicate soil functioning. We did so in a large grassland experiment which tested the effects, and relative importance, of nitrogen enrichment, plant functional composition and diversity, and foliar pathogen presence (controlled by fungicide) on soil functioning. We measured the activity of the two enzymes across seasons and years to assess the stability and temporal dynamics of soil functioning. Overall β‐glucosidase activity was slightly increased by nitrogen enrichment over time but did not respond to the other experimental treatments. Conversely, plant functional diversity and interactions between plant attributes and fungicide application were important drivers of mean acid phosphatase activity. The temporal stability of both soil enzymes was differently affected by two facets of plant diversity: species richness increased temporal stability and functional diversity decreased it; however, these effects were dampened when nitrogen and fungicide were added. The fungicide effects on soil enzyme activities suggest that foliar pathogens can also affect below‐ground processes and the interacting effect of fungicide and plant diversity suggests that these plant enemies can modulate the relationship between plant diversity and ecosystem functioning. The contrasting effects of our treatments on the mean versus stability of soil enzyme activities clearly show the need to consider temporal dynamics in below‐ground processes, to better understand the responses of soil microbes to environmental changes such as nutrient enrichment. Read the free Plain Language Summary for this article on the Journal blog.
- Open Access
- 6 July 2022
- Published by The Royal Society in Royal Society Open Science
- Vol. 9 (7) , 211283
Northward range expansion of rooting ungulates decreases detritivore and predatory mite abundances in boreal forests
In the last few decades wild boar populations have expanded northwards, colonizing boreal forests. The soil disturbances caused by wild boar rooting may have an impact on soil organisms that play a key role in organic matter turnover. However, the impact of wild boar colonization on boreal forest ecosystems and soil organisms remains largely unknown. We investigated the effect of natural and simulated rooting on decomposer and predatory soil mites (total, adult and juvenile abundances; and adult–juvenile proportion). Our simulated rooting experiment aimed to disentangle the effects of (i) bioturbation due to soil mixing and (ii) removing organic material (wild boar food resources) on soil mites. Our results showed a decline in the abundance of adult soil mites in response to both natural and artificial rooting, while juvenile abundance and the relative proportion of adults and juveniles were not affected. The expansion of wild boar northwards and into new habitats has negative effects on soil decomposer abundances in boreal forests which may cascade through the soil food web ultimately affecting ecosystem processes. Our study also suggests that a combined use of natural and controlled experimental approaches is the way forward to reveal any subtle interaction between aboveground and belowground organisms and the ecosystem functions they drive.
- 6 July 2022
- Published by The Royal Society in Royal Society Open Science
- Vol. 9 (7) , 211283
Open AccessNorthward range expansion of rooting ungulates decreases detritivore and predatory mite abundances in boreal forests
In the last few decades wild boar populations have expanded northwards, colonizing boreal forests. The soil disturbances caused by wild boar rooting may have an impact on soil organisms that play a key role in organic matter turnover. However, the impact of wild boar colonization on boreal forest ecosystems and soil organisms remains largely unknown. We investigated the effect of natural and simulated rooting on decomposer and predatory soil mites (total, adult and juvenile abundances; and adult–juvenile proportion). Our simulated rooting experiment aimed to disentangle the effects of (i) bioturbation due to soil mixing and (ii) removing organic material (wild boar food resources) on soil mites. Our results showed a decline in the abundance of adult soil mites in response to both natural and artificial rooting, while juvenile abundance and the relative proportion of adults and juveniles were not affected. The expansion of wild boar northwards and into new habitats has negative effects on soil decomposer abundances in boreal forests which may cascade through the soil food web ultimately affecting ecosystem processes. Our study also suggests that a combined use of natural and controlled experimental approaches is the way forward to reveal any subtle interaction between aboveground and belowground organisms and the ecosystem functions they drive.
- Open Access
- 22 June 2022
- Published by Wiley in GCB Bioenergy
- Vol. 14 (8) , 972-988
No evidence that conifer biochar impacts soil functioning by serving as microbial refugia in boreal soils
It is well established that application of biochar to soils can promote soil fertility, which ultimately may enhance plant growth. While many mechanisms have been proposed to explain this, one specific mechanism, the “microbial refugia hypothesis,” suggests that biochar may provide physical protection for soil microbe from soil microfauna that otherwise exert top‐down control on microbial biomass and activity. We tested the microbial refugia hypothesis by incubating two boreal soils with and without biochar derived from a wood mixture of boreal tree species (Picea abies and Pinus sylvestris), and with and without soil nematodes. We measured phospholipid fatty acids (PLFA) as a relative measure of microbial biomass, and several variables indicative of microbial activity, including extractable nutrient concentrations (NH4+, NO3−, and PO4−), heterotrophic N2‐fixation, and soil respiration. Contrary to our expectations, we found that biochar by itself did not stimulate microbial biomass or activity. Furthermore, we found that nematode addition to soil stimulated rather than depressed the biomass of several bacterial PLFA groups. Finally, interactive effects between the nematode treatment and biochar never worked in a way that supported the microbial refugia hypothesis. Our findings suggest that a typical boreal biochar applied to boreal soils may not have the same stimulatory effect on microbial biomass and activity that has been shown in some other ecosystems, and that enhanced plant growth in response to biochar addition sometimes observed in boreal environments is likely due to other mechanisms, such as direct nutrient supply from biochar or amelioration of soil pH.
- 22 June 2022
- Published by Wiley in GCB Bioenergy
- Vol. 14 (8) , 972-988
Open AccessNo evidence that conifer biochar impacts soil functioning by serving as microbial refugia in boreal soils
It is well established that application of biochar to soils can promote soil fertility, which ultimately may enhance plant growth. While many mechanisms have been proposed to explain this, one specific mechanism, the “microbial refugia hypothesis,” suggests that biochar may provide physical protection for soil microbe from soil microfauna that otherwise exert top‐down control on microbial biomass and activity. We tested the microbial refugia hypothesis by incubating two boreal soils with and without biochar derived from a wood mixture of boreal tree species (Picea abies and Pinus sylvestris), and with and without soil nematodes. We measured phospholipid fatty acids (PLFA) as a relative measure of microbial biomass, and several variables indicative of microbial activity, including extractable nutrient concentrations (NH4+, NO3−, and PO4−), heterotrophic N2‐fixation, and soil respiration. Contrary to our expectations, we found that biochar by itself did not stimulate microbial biomass or activity. Furthermore, we found that nematode addition to soil stimulated rather than depressed the biomass of several bacterial PLFA groups. Finally, interactive effects between the nematode treatment and biochar never worked in a way that supported the microbial refugia hypothesis. Our findings suggest that a typical boreal biochar applied to boreal soils may not have the same stimulatory effect on microbial biomass and activity that has been shown in some other ecosystems, and that enhanced plant growth in response to biochar addition sometimes observed in boreal environments is likely due to other mechanisms, such as direct nutrient supply from biochar or amelioration of soil pH.
- Open Access
- 20 February 2020
- Published by Frontiers Media SA in Frontiers in Forests and Global Change
- Vol. 3
Global Change Impacts on Forest Soils: Linkage Between Soil Biota and Carbon-Nitrogen-Phosphorus Stoichiometry
Forest ecosystems are subjected to global change drivers worldwide, such as increasing temperature, atmospheric carbon dioxide, nutrient pollution, as well as changes in fire and precipitation regimes. These global change drivers have greatly modified the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), which has an impact on primary productivity in forest ecosystems and in turn, affect the quality and quantity of resources entering the soil food web. However, C, N, and P soil dynamics have been mostly studied without considering their coupling effects on soil organisms. This is of critical interest because changes in nutrient stoichiometry may have a strong effect on soil biota and the ecosystem functions they drive. Further, most studies have focused on global change effects on bacteria and fungi and their C:N:P stoichiometry, while neglecting other soil organisms at higher trophic levels. This has led to an incomplete understanding of how the entire soil food web drives ecosystem processes involved in organic matter turnover and nutrient cycling. Here, we review studies that investigated how global change drivers impact C:N:P stoichiometry of soil organisms at different trophic levels in forest ecosystems and identify important knowledge gaps. We propose future directions for research on global change impacts on the linkages between soil biota and C:N:P stoichiometry.
- 20 February 2020
- Published by Frontiers Media SA in Frontiers in Forests and Global Change
- Vol. 3
Open AccessGlobal Change Impacts on Forest Soils: Linkage Between Soil Biota and Carbon-Nitrogen-Phosphorus Stoichiometry
Forest ecosystems are subjected to global change drivers worldwide, such as increasing temperature, atmospheric carbon dioxide, nutrient pollution, as well as changes in fire and precipitation regimes. These global change drivers have greatly modified the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), which has an impact on primary productivity in forest ecosystems and in turn, affect the quality and quantity of resources entering the soil food web. However, C, N, and P soil dynamics have been mostly studied without considering their coupling effects on soil organisms. This is of critical interest because changes in nutrient stoichiometry may have a strong effect on soil biota and the ecosystem functions they drive. Further, most studies have focused on global change effects on bacteria and fungi and their C:N:P stoichiometry, while neglecting other soil organisms at higher trophic levels. This has led to an incomplete understanding of how the entire soil food web drives ecosystem processes involved in organic matter turnover and nutrient cycling. Here, we review studies that investigated how global change drivers impact C:N:P stoichiometry of soil organisms at different trophic levels in forest ecosystems and identify important knowledge gaps. We propose future directions for research on global change impacts on the linkages between soil biota and C:N:P stoichiometry.
- Open Access
- 23 January 2020
- Published by Springer Nature in Ecosystems
- Vol. 23 (7) , 1423-1436
Low and High Nitrogen Deposition Rates in Northern Coniferous Forests Have Different Impacts on Aboveground Litter Production, Soil Respiration, and Soil Carbon Stocks
Nitrogen (N) deposition can change the carbon (C) sink of northern coniferous forests by changing the balance between net primary production and soil respiration. We used a field experiment in an N poor Pinus sylvestris forest where five levels of N (0, 3, 6, 12, and 50 kg N ha−1 yr−1, n = 6) had been added annually for 12–13 years to investigate how litter C inputs and soil respiration, divided into its autotrophic and heterotrophic sources, respond to different rates of N input, and its subsequent effect on soil C storage. The highest N addition rate (50 kg N ha−1 yr−1) stimulated soil C accumulation in the organic layer by 22.3 kg C kg−1 N added, increased litter inputs by 46%, and decreased soil respiration per mass unit of soil C by 31.2%, mainly by decreasing autotrophic respiration. Lower N addition rates (≤ 12 kg N ha−1 yr−1) had no effect on litter inputs or soil respiration. These results support previous studies reporting on increased litter inputs coupled to impeded soil C mineralization, contributing to enhancing the soil C sink when N is supplied at high rates, but add observations for lower N addition rates more realistic for N deposition. In doing so, we show that litter production in N poor northern coniferous forests can be relatively unresponsive to low N deposition levels, that stimulation of microbial activity at low N additions is unlikely to reduce the soil C sink, and that high levels of N deposition enhance the soil C sink by increasing litter inputs and decreasing soil respiration.
- 23 January 2020
- Published by Springer Nature in Ecosystems
- Vol. 23 (7) , 1423-1436
Open AccessLow and High Nitrogen Deposition Rates in Northern Coniferous Forests Have Different Impacts on Aboveground Litter Production, Soil Respiration, and Soil Carbon Stocks
Nitrogen (N) deposition can change the carbon (C) sink of northern coniferous forests by changing the balance between net primary production and soil respiration. We used a field experiment in an N poor Pinus sylvestris forest where five levels of N (0, 3, 6, 12, and 50 kg N ha−1 yr−1, n = 6) had been added annually for 12–13 years to investigate how litter C inputs and soil respiration, divided into its autotrophic and heterotrophic sources, respond to different rates of N input, and its subsequent effect on soil C storage. The highest N addition rate (50 kg N ha−1 yr−1) stimulated soil C accumulation in the organic layer by 22.3 kg C kg−1 N added, increased litter inputs by 46%, and decreased soil respiration per mass unit of soil C by 31.2%, mainly by decreasing autotrophic respiration. Lower N addition rates (≤ 12 kg N ha−1 yr−1) had no effect on litter inputs or soil respiration. These results support previous studies reporting on increased litter inputs coupled to impeded soil C mineralization, contributing to enhancing the soil C sink when N is supplied at high rates, but add observations for lower N addition rates more realistic for N deposition. In doing so, we show that litter production in N poor northern coniferous forests can be relatively unresponsive to low N deposition levels, that stimulation of microbial activity at low N additions is unlikely to reduce the soil C sink, and that high levels of N deposition enhance the soil C sink by increasing litter inputs and decreasing soil respiration.