the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Unraveling biogeographical patterns and environmental drivers of soil fungal diversity at the French national scale
Christophe Djemiel
Samuel Dequiedt
Walid Horrigue
Arthur Bailly
Mélanie Lelièvre
Julie Tripied
Charles Guilland
Solène Perrin
Gwendoline Comment
Nicolas Saby
Claudy Jolivet
Antonio Bispo
Line Boulonne
Antoine Pierart
Patrick Wincker
Corinne Cruaud
Pierre-Alain Maron
Sébastien Terrat
Lionel Ranjard
Abstract. The fungal kingdom is among the most diversified kingdoms on earth with estimations up to 12 million species. Yet, it remains poorly understood with only 150,000 fungal species currently described. Given the major ecological role of fungi in ecosystem functioning, these numbers stress the importance of investigating fungal diversity description across different ecosystem types. Here, we explored the spatial distribution of the soil fungal diversity on a broad geographical scale, using the French Soil Quality Monitoring Network that covers the whole French territory (2,171 soils sampled along a systematic grid). Fungal alpha-diversity was assessed directly from soil DNA using a metabarcoding approach. Total cumulated fungal diversity across France included 136,219 OTUs, i.e., about 1 % of the global soil fungal diversity for a territory representing only 0.3 % of terrestrial surface on Earth. Based on this dataset, the first extensive map of fungal alpha-diversity was drawn and evidenced a heterogeneous and spatially structured distribution in large biogeographical patterns of 231 km radius for richness (Hill number q=0) and smaller patterns of 36 km radius for dominant fungi (Hill number q=2). As related to other environmental parameters, the spatial distribution of fungal diversity was mainly influenced by local filters such as soil characteristics and land management, but also by global filters such as climate conditions. The spatial distribution of abundant and rare fungi was determined by distinct or similar filters with various relative influences. Interestingly, cropped soils exhibited the highest pool of fungal diversity relatively to forest and vineyard soils. In complement, soil fungal OTUs network interactions were calculated under the different land uses across France. They varied hugely and showed a loss of 75 % of the complexity in crop systems and grasslands compared to forests, and up to 83 % in vineyard systems. Overall, our study revealed that a nation-wide survey with a high spatial resolution approach is relevant to deeply investigate the spatial distribution and determinism of soil fungal diversity. Our findings provide novel insights for a better understanding of soil fungal ecology and upgrade biodiversity conservation policies by supplying representative repositories dedicated to soil microorganisms in a context of global change.
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Christophe Djemiel et al.
Status: open (until 01 Nov 2023)
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RC1: 'Comment on egusphere-2023-1552', César Marín, 10 Sep 2023
reply
The manuscript entitled “Unraveling biogeographical patterns and environmental drivers of soil fungal diversity at the French national scale”, by Djemiel and co-authors investigated soil fungal diversity across a vast sampling in France. The article is well written and analyzed, and makes use of a huge amount of data… sadly, the selected molecular marker is not good/precise enough to elucidate fungal diversity.
Abstract
L.40. Change “approach. Total cumulated” to “approach. The total accumulated”.
L.41. This “1%” is estimated based on which number? In a number mentioned above (ie. 12 million species), or on what? Try to be a bit more specific.
L.49. Here, “distinct or similar filters” relative to what?
Introduction
L.80. Not only nitrogen but also phosphorous.
L.87. Change “agents and involved” to “agents and are involved”.
L.94-97. This is sort of true… but, check more recent, global sampling by the same author (Leho Tedersoo), as DOI: 10.1007/s13225-021-00493-7 and DOI: 10.1111/gcb.16398, among others.
L.130. Change “altitude) and climate” to “altitude), and climate”.
L.140. Change “types and soil characteristics” to “types, and soil characteristics of the country”.
L.141. How do you define pedological diversity?
L.142. Change “(Minasny et al., 2010) and is also” to “(Minasny et al., 2010), and is also”.
L.144. Change “grasslands and forests” to “grasslands, and forests”.
L.146. Change “statements” to “facts”.
L.146-148. This is a very vague phrase… as a researcher based in Chile, I can say we also say this all the time… all countries are unique, thus, this phrase is not.
L.149. Change “sampling strategy in the world and the systematic random” to “sampling strategies in the world, and this systematic random”.
L.154. Change “bioinformatics and statistical” to “bioinformatics, and statistical”.
L.156-157. At this point I have not read the rest of the manuscript… but it does make no sense at all to sequence the 18S to characterize soil fungal taxa. ITS1 or ITS2 are the regions to be sequenced.
L.163. Change “diversity and inverse” to “diversity, and inverse”. Change “prediction, (geostatistics)” to “prediction (geostatistics)”.
L.166. Change “conditions and land use” to “conditions, and land use”.
L.168. I do not get what you mean by “In fine”.
Methods
L.178. Change “(n=65) and low anthropized” to “(n=65), and low anthropized”.
L.181. Change “were bulked” to “were combined”.
L.183. For how long these samples were air-dried? When too long, a bunch of fungal taxa might be lost. The correct approach is: either to dry the soil samples in oven (max. 40 C) or with silica gel, or freeze the samples (at -20 C or -80 C) less than 24 hours after collecting.
L.185-187. Still, some brief description of soil chemical analyses is needed here.
L.195. Change “beads and 4 glass beads” to “beads, and 4 glass beads”.
L.197. Change “sulfate and up to” to “sulfate, and up to”.
L.209-211. Sadly, this selected regions is VERY inefficient to detect fungal diversity, ie. it only captures a very small fraction. It has happened to me.
L.245. When you say “using the shapiro.test function”, I know you refer to R, but you need to write of what package is that function (and correctly cite such package), and also, on which software -R- and cite such software.
L.247. Change “from forecast package” to “from the R forecast package”.
L.257. Change “as recommended by (Dini‐Andreote et al., 2021).” to “as recommended by Dini‐Andreote et al. (2021).”, but, I am not sure what you mean by this.
L.266. Change “the “usdm” package” to “the “usdm” R package”.
L.268. Change ““leaps” package” to ““leaps” R package”.
L.271. Change “functions (“vegan” package (Oksanen et al., 2013)).” to “functions of the “vegan” R package Oksanen et al. (2013)).”.
L.272. Change “forward selection to” to “forward multiple regression selection to”.
L.277. Change “in (Granger et al., 2015).” to “in Granger et al. (2015).”.
L.279. Here, “using a local neighborhood” what?
L.280. Change “the “gstat” package” to “the “gstat” R package”.
L.306. Cite this software correctly.
Results
L.328. Change “south-east and the” to “south-east, and the”.
L.340. Change “conditions and spatial” to “conditions, and spatial”.
L.345. Change “(0.39%) and spatial” to “(0.39%), and spatial”.
L.347. Change “(0.63%) and land use” to “(0.63%), and land use”.
L.350. Change “for q=1 and neared” to “for q=1, and neared”.
L.355. Change “was the pH” to “was pH”.
L.361, 363. Change “the soil pH” to “soil pH”.
L.364. Here, “were also identified” as what?
L.379. Change “soils and can be ranked” to “soils, and can be ranked”.
L.405. Change “with forest” to “with a pattern: forest”.
Discussion
L.420. Again, “1%” relative to what?
L.427. Change “were collected sites on” to “were collected in sites on”.
L.441. Change “hence possible biases” to “hence with possible biases”.
L.454. Change “OTUs), “typical”” to “OTUs), and “typical””.
L.457. Change “radius and “spottier”” to “radius, and “spottier””.
L.469. Change “the soil pH” to “soil pH”. Change “by the clay content” to “by clay content”.
L.473. Change “of the pH in the” to “of soil pH in the”.
L.475. Change “the soil pH” to “soil pH”.
L.496-498. The last part of this phrase (“whatever the Hill number”) is not clear.
L.516. Change “2017) and fungi” to “2017), and fungi”.
L.517. Change “grassland and crop” to “grassland, and crop”.
L.534. Change “soil structure and biological” to “soil structure, and biological”.
Conclusions
L.584. Change “processes and environmental” to “processes, and environmental”.
Citation: https://doi.org/10.5194/egusphere-2023-1552-RC1 -
AC1: 'Reply on RC1', Christophe Djemiel, 15 Sep 2023
reply
Reviewer
The manuscript entitled “Unraveling biogeographical patterns and environmental drivers of soil fungal diversity at the French national scale”, by Djemiel and co-authors investigated soil fungal diversity across a vast sampling in France. The article is well written and analyzed, and makes use of a huge amount of data… sadly, the selected molecular marker is not good/precise enough to elucidate fungal diversity.
Author comment on behalf of all Co-Authors (AC) to reviewer:
We agree with the reviewer that “ITS” is the most common and historical molecular marker for assessing fungal taxonomy, especially to describe taxonomic groups. However, 18S rDNA has also been an alternative in the last few years to assess the soil fungal community structure and richness more globally, as demonstrated by several authors – in particular in biogeography and at large scales (see below).
- George, P. B., Creer, S., Griffiths, R. I., Emmett, B. A., Robinson, D. A., & Jones, D. L. (2019). Primer and database choice affect fungal functional but not biological diversity findings in a national soil survey. Frontiers in Environmental Science, 173.
- Li, J., Delgado-Baquerizo, M., Wang, J. T., Hu, H. W., Cai, Z. J., Zhu, Y. N., & Singh, B. K. (2019). Fungal richness contributes to multifunctionality in boreal forest soil. Soil Biology and Biochemistry, 136, 107526.
- Bastida, F., Eldridge, D. J., García, C., Kenny Png, G., Bardgett, R. D., & Delgado-Baquerizo, M. (2021). Soil microbial diversity–biomass relationships are driven by soil carbon content across global biomes. The ISME Journal, 15(7), 2081-2091.
- Fernandes, M. L. P., Bastida, F., Jehmlich, N., Martinović, T., Větrovský, T., Baldrian, P., ... & Starke, R. (2022). Functional soil mycobiome across ecosystems. Journal of Proteomics, 252, 104428.
We will add a specific paragraph in the discussion of the manuscript justifying the relevance of our molecular marker, and more broadly about the metabarcoding approach (between L432 and L433).
Relevance and drawbacks of the metabarcoding approach
Two molecular markers are commonly used to explore fungal diversity thanks to metabarcoding approaches: the internal transcribed spacer (ITS) region, accepted as a universal barcode, and the 18S rRNA gene as an alternative. Both have advantages and drawbacks, in particular to observe specific functional groups. For example, members of the class Glomeromycetes are better characterized using 18S rDNA than ITS, especially in the soil microbiota, and this could have a significant impact on fungal diversity metrics (George et al., 2019). In addition, important reference sequences are only annotated at the phylum level in the international databases (Nilsson et al., 2012; Nilsson et al., 2016; Banos et al., 2018). Moreover, a recent study highlighted that national-scale fungal biogeography studies based on 18S rDNA were relevant to decipher the relationships between fungal diversity and environmental filters (George et al., 2019). For all these reasons, we chose to use the 18S rDNA gene to characterize fungal diversity. Once the molecular marker was chosen, the hypervariable region had to be selected for sequencing. Various criteria had to be taken into account such as amplicon length in relation to the sequencer, or the desired taxonomic and phylogenetic resolution. We sequenced the V7-V8 regions because they appeared to be the most promising regions for fungal diversity assessment (Banos et al., 2018). The last tricky step of a fungal diversity study is the bioinformatics analysis, which is dependent on the pipeline (Pauvert et al., 2019). Nevertheless, our previous studies on bacterial biogeography showed that our metabarcoding pipeline and associated tools were highly appropriate to analyze large microbial datasets (Djemiel et al., 2020; Terrat et al., 2019).Although alpha-diversity depends on the intensity and strategy of sampling, especially when addressing the complexity of the soil microbiota (Willis et al., 2019; Castle et al., 2019), it is valuable in microbial ecology. As for the alpha-diversity analysis, we chose to use the Hill numbers – which have several advantages (Roswell et al., 2021) – in order to provide an overview of fungal diversity within sites. This allowed us to observe all OTUs, "typical" OTUs and dominant ones, based on their abundance frequencies (Chao et al., 2014), especially as fungal diversity can be represented by a few dominant species with a high relative abundance (Egidi et al., 2019).
- George, P. B., Creer, S., Griffiths, R. I., Emmett, B. A., Robinson, D. A., & Jones, D. L. (2019). Primer and database choice affect fungal functional but not biological diversity findings in a national soil survey. Frontiers in Environmental Science, 173.
- Nilsson, R. H., Tedersoo, L., Abarenkov, K., Ryberg, M., Kristiansson, E., Hartmann, M., ... & Kõljalg, U. (2012). Five simple guidelines for establishing basic authenticity and reliability of newly generated fungal ITS sequences. MycoKeys, 4, 37-63.
- Nilsson, R. H., Wurzbacher, C., Bahram, M., Coimbra, V. R., Larsson, E., Tedersoo, L., ... & Abarenkov, K. (2016). Top 50 most wanted fungi. MycoKeys, (12), 29-40.
- Banos, S., Lentendu, G., Kopf, A., Wubet, T., Glöckner, F. O., & Reich, M. (2018). A comprehensive fungi-specific 18S rRNA gene sequence primer toolkit suited for diverse research issues and sequencing platforms. BMC microbiology, 18, 1-15.
- Pauvert, C., Buée, M., Laval, V., Edel-Hermann, V., Fauchery, L., Gautier, A., ... & Vacher, C. (2019). Bioinformatics matters: The accuracy of plant and soil fungal community data is highly dependent on the metabarcoding pipeline. Fungal Ecology, 41, 23-33.
- Djemiel, C., Dequiedt, S., Karimi, B., Cottin, A., Girier, T., El Djoudi, Y., ... & Terrat, S. (2020). BIOCOM-PIPE: a new user-friendly metabarcoding pipeline for the characterization of microbial diversity from 16S, 18S and 23S rRNA gene amplicons. BMC bioinformatics, 21, 1-21.
- Terrat, S., Djemiel, C., Journay, C., Karimi, B., Dequiedt, S., Horrigue, W., ... & Ranjard, L. (2020). ReClustOR: a re‐clustering tool using an open‐reference method that improves operational taxonomic unit definition. Methods in Ecology and Evolution, 11(1), 168-180.
- Willis, A. D. (2019). Rarefaction, alpha diversity, and statistics. Frontiers in microbiology, 10, 2407.
- Castle, S. C., Samac, D. A., Sadowsky, M. J., Rosen, C. J., Gutknecht, J. L., & Kinkel, L. L. (2019). Impacts of sampling design on estimates of microbial community diversity and composition in agricultural soils. Microbial ecology, 78, 753-763.
- Roswell, M., Dushoff, J., & Winfree, R. (2021). A conceptual guide to measuring species diversity. Oikos, 130(3), 321-338.
- Chao, A., Gotelli, N. J., Hsieh, T. C., Sander, E. L., Ma, K. H., Colwell, R. K., & Ellison, A. M. (2014). Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecological monographs, 84(1), 45-67.
- Egidi, E., Delgado-Baquerizo, M., Plett, J. M., Wang, J., Eldridge, D. J., Bardgett, R. D., ... & Singh, B. K. (2019). A few Ascomycota taxa dominate soil fungal communities worldwide. Nature communications, 10(1), 2369.
Abstract
L.40. Change “approach. Total cumulated” to “approach. The total accumulated”.
We agree, we are going to change the sentence.
L.41. This “1%” is estimated based on which number? In a number mentioned above (ie. 12 million species), or on what? Try to be a bit more specific.
Yes indeed, we based ourselves on the most recent estimation of fungal diversity (Phukhamsakda et al., 2022). We completed the sentence.
“Total cumulated fungal diversity across France included 136,219 OTUs, i.e., about 1% of the global soil fungal diversity (based on a maximum diversity estimate of 12 million) for a territory representing only 0.3% of terrestrial surface on Earth.”
L.49. Here, “distinct or similar filters” relative to what?
We agree, we completed the sentence by adding the distinct and similar drivers of the Hill numbers q=0 and q=2).
The spatial distribution of abundant (q=2) and rare (q=0) fungi was determined by distinct (clay or organic carbon) or similar (soil pH) filters with various relative influences.
Introduction
L.80. Not only nitrogen but also phosphorous.
We fully agree, thanks for pointing this out, we added phosphorous in the sentence.
The functions of fungal communities provide many ecosystem services that promote mineral nutrition of plants linked to soil organic matter rates, phosphorous and nitrogen availability (Miyauchi et al., 2020; Ward et al., 2022).
L.87. Change “agents and involved” to “agents and are involved”.
We agree, we are going to change the sentence.
L.94-97. This is sort of true… but, check more recent, global sampling by the same author (Leho Tedersoo), as DOI: 10.1007/s13225-021-00493-7 and DOI: 10.1111/gcb.16398, among others.
We agree, we are going to add this reference study in the sentence.
L.130. Change “altitude) and climate” to “altitude), and climate”.
We agree, we are going to change the sentence.
L.140. Change “types and soil characteristics” to “types, and soil characteristics of the country”.
We agree, we are going to change the sentence.
L.141. How do you define pedological diversity?
We defined pedological diversity as the number of different soil types. The complete definition from the publication reads as follows, “The conventional pedodiversity measure is related to the abundance of the observed soil type, thus there is a relationship between land area and Shannon's entropy.” (Minasny, Budiman, Alex B. McBratney, and Alfred E. Hartemink. "Global pedodiversity, taxonomic distance, and the World Reference Base." Geoderma 155.3-4 (2010): 132-139.)
L.142. Change “(Minasny et al., 2010) and is also” to “(Minasny et al., 2010), and is also”.
We agree, we are going to change the sentence.
L.144. Change “grasslands and forests” to “grasslands, and forests”.
We agree, we are going to change the sentence.
L.146. Change “statements” to “facts”.
We agree, we are going to change the sentence.
L.146-148. This is a very vague phrase… as a researcher based in Chile, I can say we also say this all the time… all countries are unique, thus, this phrase is not.
We fully agree, all countries are unique but the wide range of variation of climates, soils and land uses in France combined with an intensive and extensive soil sampling strategy leads us to conclude about the relevance of our study.
See our two sentences L146 to L151
L.149. Change “sampling strategy in the world and the systematic random” to “sampling strategies in the world, and this systematic random”.
We agree, we are going to change the sentence.
L.154. Change “bioinformatics and statistical” to “bioinformatics, and statistical”.
We agree, we are going to change the sentence.
L.156-157. At this point I have not read the rest of the manuscript… but it does make no sense at all to sequence the 18S to characterize soil fungal taxa. ITS1 or ITS2 are the regions to be sequenced.
We have detailed this response above, see the overall comment and the text that will be added to our discussion.
L.163. Change “diversity and inverse” to “diversity, and inverse”. Change “prediction, (geostatistics)” to “prediction (geostatistics)”.
We agree, we are going to change the sentence.
L.166. Change “conditions and land use” to “conditions, and land use”.
We agree, we are going to change the sentence.
L.168. I do not get what you mean by “In fine”.
Borrowed from Latin in fine, but we can change to “Finally”.
Methods
L.178. Change “(n=65) and low anthropized” to “(n=65), and low anthropized”.
We agree, we are going to change the sentence.
L.181. Change “were bulked” to “were combined”.
We agree, we are going to change the sentence.
L.183. For how long these samples were air-dried? When too long, a bunch of fungal taxa might be lost. The correct approach is: either to dry the soil samples in oven (max. 40 C) or with silica gel, or freeze the samples (at -20 C or -80 C) less than 24 hours after collecting.
We agree, we will specify this point in the text.
Each sample was air-dried following a standardized procedure at 35 °C until the soil humidity was below 1%, then sieved to 2 mm and separated into two sub-samples.
L.185-187. Still, some brief description of soil chemical analyses is needed here.
We agree, we are going to change the sentence.
A detailed description of the physicochemical analysis performed in this study (soil pH, texture, organic carbon, nitrogen, and phosphorus) is accessible from (Jolivet et al., 2006).
L.195. Change “beads and 4 glass beads” to “beads, and 4 glass beads”.
We agree, we are going to change the sentence.
L.197. Change “sulfate and up to” to “sulfate, and up to”.
We agree, we are going to change the sentence.
L.209-211. Sadly, this selected regions is VERY inefficient to detect fungal diversity, ie. it only captures a very small fraction. It has happened to me.
We have detailed this response above, see the overall comment and the text that will be added to our discussion.
In addition, Banos et al. (2018) indicate that the regions we chose are most robust for characterizing fungal communities.
L.245. When you say “using the shapiro.test function”, I know you refer to R, but you need to write of what package is that function (and correctly cite such package), and also, on which software -R- and cite such software.
We agree, we are going to clarify the sentence.
We tested if the variables were normally distributed or approximately so, using the shapiro.test function (R “stats” package).
L.247. Change “from forecast package” to “from the R forecast package”.
We agree, we are going to change the sentence.
L.257. Change “as recommended by (Dini‐Andreote et al., 2021).” to “as recommended by Dini‐Andreote et al. (2021).”, but, I am not sure what you mean by this.
We agree, we are going to clarify the sentence.
The details of the samples removed (e.g., outliers) in the different land uses are available in Supplementary Figure 1, as recommended by Dini‐Andreote et al. (2021) (Dini‐Andreote et al., 2021), who developed a data management strategy with good practices for biogeographical studies.
L.266. Change “the “usdm” package” to “the “usdm” R package”.
We agree, we are going to change the sentence.
L.268. Change ““leaps” package” to ““leaps” R package”.
We agree, we are going to change the sentence.
L.271. Change “functions (“vegan” package (Oksanen et al., 2013)).” to “functions of the “vegan” R package Oksanen et al. (2013)).”.
We agree, we are going to change the sentence.
L.272. Change “forward selection to” to “forward multiple regression selection to”.
We agree, we are going to change the sentence.
L.277. Change “in (Granger et al., 2015).” to “in Granger et al. (2015).”.
We agree, we are going to change the sentence.
L.279. Here, “using a local neighborhood” what?
Yes, we replaced the sentence by the following one:
Then, we predicted the unsampled locations by a global kriging approach that used all the points in the dataset (global neighborhood).
L.280. Change “the “gstat” package” to “the “gstat” R package”.
We agree, we are going to change the sentence.
L.306. Cite this software correctly.
We agree, we are going to change the sentence.
The networks were mapped using Cytoscape software (v. 3.9.1) (Shannon et al., 2003).
Shannon, P., Markiel, A., Ozier, O., Baliga, N. S., Wang, J. T., Ramage, D., ... & Ideker, T. (2003). Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome research, 13(11), 2498-2504.
Results
L.328. Change “south-east and the” to “south-east, and the”.
We agree, we are going to change the sentence.
L.340. Change “conditions and spatial” to “conditions, and spatial”.
We agree, we are going to change the sentence.
L.345. Change “(0.39%) and spatial” to “(0.39%), and spatial”.
We agree, we are going to change the sentence.
L.347. Change “(0.63%) and land use” to “(0.63%), and land use”.
We agree, we are going to change the sentence.
L.350. Change “for q=1 and neared” to “for q=1, and neared”.
We agree, we are going to change the sentence.
L.355. Change “was the pH” to “was pH”.
We agree, we are going to change the sentence.
L.361, 363. Change “the soil pH” to “soil pH”.
We agree, we are going to change the sentence.
L.364. Here, “were also identified” as what?
We agree, we are going to change the sentence.
The soil pH remained the strongest driver with also a unimodal relationship, and organic carbon and the total iron content were also identified as secondary drivers, with a negative linear relationship.
L.379. Change “soils and can be ranked” to “soils, and can be ranked”.
We agree, we are going to change the sentence.
L.405. Change “with forest” to “with a pattern: forest”.
We agree, we are going to change the sentence.
Discussion
L.420. Again, “1%” relative to what?
We agree, we are going to change the sentence.
Compared to the estimated worldwide diversity, France exhibits a very high cumulated soil fungal richness (about 1% of the global soil fungal diversity based on a maximum diversity estimate of 12 million) relative to its small surface (0.3 % of terrestrial land).
L.427. Change “were collected sites on” to “were collected in sites on”.
We agree, we are going to change the sentence.
L.441. Change “hence possible biases” to “hence with possible biases”.
We agree, we are going to change the sentence.
L.454. Change “OTUs), “typical”” to “OTUs), and “typical””.
We agree, we are going to change the sentence.
L.457. Change “radius and “spottier”” to “radius, and “spottier””.
We agree, we are going to change the sentence.
L.469. Change “the soil pH” to “soil pH”. Change “by the clay content” to “by clay content”.
We agree, we are going to change the sentence.
L.473. Change “of the pH in the” to “of soil pH in the”.
We agree, we are going to change the sentence.
L.475. Change “the soil pH” to “soil pH”.
We agree, we are going to change the sentence.
L.496-498. The last part of this phrase (“whatever the Hill number”) is not clear.
We removed the « whatever the Hill number (Terrat et al., 2017). To clarify the text
L.516. Change “2017) and fungi” to “2017), and fungi”.
We agree, we are going to change the sentence.
L.517. Change “grassland and crop” to “grassland, and crop”.
We agree, we are going to change the sentence.
L.534. Change “soil structure and biological” to “soil structure, and biological”.
We agree, we are going to change the sentence.
Conclusions
L.584. Change “processes and environmental” to “processes, and environmental”.
We agree, we are going to change the sentence.
Citation: https://doi.org/10.5194/egusphere-2023-1552-AC1
-
AC1: 'Reply on RC1', Christophe Djemiel, 15 Sep 2023
reply
-
EC1: 'Comment on egusphere-2023-1552', Elizabeth Bach, 25 Sep 2023
reply
Apologies for how slow this process is proceeding; I'm having a very difficult time finding a second reviewer. I've nominated additional reviewers and am hoping for a positive response soon.
Citation: https://doi.org/10.5194/egusphere-2023-1552-EC1
Christophe Djemiel et al.
Christophe Djemiel et al.
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