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The global contribution of soil mosses to ecosystem services


Soil mosses are among the most widely distributed organisms on land. Experiments and observations suggest that they contribute to terrestrial soil biodiversity and function, yet their ecological contribution to soil has never been assessed globally under natural conditions. Here we conducted the most comprehensive global standardized field study to quantify how soil mosses influence 8 ecosystem services associated with 24 soil biodiversity and functional attributes across wide environmental gradients from all continents. We found that soil mosses are associated with greater carbon sequestration, pool sizes for key nutrients and organic matter decomposition rates but a lower proportion of soil-borne plant pathogens than unvegetated soils. Mosses are especially important for supporting multiple ecosystem services where vascular-plant cover is low. Globally, soil mosses potentially support 6.43 Gt more carbon in the soil layer than do bare soils. The amount of soil carbon associated with mosses is up to six times the annual global carbon emissions from any altered land use globally. The largest positive contribution of mosses to soils occurs under perennial, mat and turf mosses, in less-productive ecosystems and on sandy soils. Our results highlight the contribution of mosses to soil life and functions and the need to conserve these important organisms to support healthy soils.

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Fig. 1: A global survey of mosses to investigate soil biodiversity and function.
Fig. 2: Global distribution of soil mosses.
Fig. 3: Contribution of mosses and vascular plants to multiple ecosystem services.
Fig. 4: Contribution of vascular plants and mosses to multiservices.
Fig. 5: Environmental factors associated with the contribution of mosses to multiple ecosystem services.

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Data availability

All the materials, raw data, and protocols used in the article are available upon request. Data used in this study can be found in the Figshare data repository ref. 51.


  1. Lindo, Z. & Gonzalez, A. The Bryosphere: an integral and influential component of the Earth’s biosphere. Ecosystems 13, 612–627 (2010).

    Article  Google Scholar 

  2. Turetsky, M. R. et al. The resilience and functional role of moss in boreal and arctic ecosystems. New Phytol. 196, 49–67 (2012).

    Article  Google Scholar 

  3. Rodriguez-Caballero, E. et al. Dryland photoautotrophic soil surface communities endangered by global change. Nat. Geosci. 11, 185–189 (2018).

    Article  Google Scholar 

  4. Shaw, A. J., Cox, C. J. & Goffinet, B. Global patterns of moss diversity: taxonomic and molecular inferences. Taxon 54, 337–352 (2005).

    Article  Google Scholar 

  5. Garcia-Moya, E. & McKell, C. M. Contribution of shrubs to the nitrogen economy of a desert-wash plant community. Ecology 51, 81–88 (1970).

    Article  Google Scholar 

  6. Jonsson, M. et al. Direct and indirect drivers of moss community structure, function, and associated microfauna across a successional gradient. Ecosystems 18, 154–169 (2015).

    Article  Google Scholar 

  7. Delgado-Baquerizo, M. et al. Biocrust forming mosses mitigate the impact of aridity on soil microbial communities in drylands: observational evidence from three continents. New Phytol. 220, 824–835 (2018).

    Article  Google Scholar 

  8. Eldridge, D. J. et al. The pervasive and multifaceted influence of biocrusts on water in the world’s drylands. Glob. Change Biol. 26, 6003–6014 (2020).

    Article  Google Scholar 

  9. Kasimir, Å., He, H., Jansson, P.-E., Lohila, A. & Minkkinen, K. Mosses are important for soil carbon sequestration in forested peatlands. Front. Environ. Sci. 9, 680430 (2021).

    Article  Google Scholar 

  10. Reed, S. C. et al. Changes to dryland rainfall result in rapid moss mortality and altered soil fertility. Nat. Clim. Change 2, 752–755 (2012).

    Article  Google Scholar 

  11. Romero, A. N., Moratta, M. H., Vento, B., Rodriguez, R. & Carretero, E. M. Variations in the coverage of biological soil crusts along an aridity gradient in the central-west Argentina. Acta Oecol. 109, 103671 (2020).

    Article  Google Scholar 

  12. Byun, M. Y., Kim, D., Youn, U. J., Lee, S. & Lee, H. Improvement of moss photosynthesis by humic acids from Antarctic tundra soil. Plant Physiol. Biochem. 159, 37–42 (2021).

    Article  Google Scholar 

  13. Gross, N. et al. Linking individual response to biotic interactions with community structure: a trait-based framework. Funct. Ecol. 23, 1167–1178 (2009).

    Article  Google Scholar 

  14. Büdel, B. et al. Improved appreciation of the functioning and importance of biological soil crusts in Europe: the Soil Crust International Project (SCIN). Biodivers. Conserv. 23, 1639–1658 (2014).

    Article  Google Scholar 

  15. Geffert, J. L., Frahn, J. L., Barthlott, W. & Mutke, J. Global moss diversity: spatial and taxonomic patterns of species richness. J. Bryol. 35, 1–11 (2013).

    Article  Google Scholar 

  16. Delgado-Baquerizo, M. et al. Multiple elements of soil biodiversity drive ecosystem functions across biomes. Nat. Ecol. Evol. 4, 210–220 (2020).

    Article  Google Scholar 

  17. Armas, C., Ordiales, R. & Pugnaire, F. I. Measuring plant interactions: a new comparative index. Ecology 85, 2682–2686 (2004).

    Article  Google Scholar 

  18. Hollingsworth, T. N., Schuur, E. A. G., Chapin, F. S. & Walker, M. D. Plant community composition as a predictor of regional soil carbon storage in Alaskan boreal black spruce ecosystems. Ecosystems 11, 629–642 (2008).

    Article  Google Scholar 

  19. Porada, P., Weber, B., Elbert, W., Pöschl, U. & Kleidon, A. Estimating global carbon uptake by lichens and bryophytes with a process-based model. Biogeosciences 10, 6989–7033 (2013).

    Article  Google Scholar 

  20. LaRoi, G. H. & Stringer, M. H. Ecological studies in the boreal spruce–fir forests of the North American taiga. II. Analysis of the bryophyte flora. Can. J. Bot. 54, 619–643 (1976).

    Article  Google Scholar 

  21. Ino, Y. & Nakatsubo, T. Distribution of carbon, nitrogen and phosphorus in a moss community soil system developed on a cold desert in Antarctica. Ecol. Res. 1, 59–69 (1986).

    Article  Google Scholar 

  22. Chapin, F. III, Oechel, W., Van Cleve, K. & Lawrence, W. The role of mosses in the phosphorus cycling of an Alaskan black spruce forest. Oecologia 74, 310–315 (1987).

    Article  Google Scholar 

  23. Brown, D. H. & Bates, J. W. Bryophytes and nutrient cycling. Bot. J. Linn. Soc. 104, 129–147 (1990).

    Article  Google Scholar 

  24. Makajanma, M. M., Taufik, I. & Faizal, A. Antioxidant and antibacterial activity of extract from two species of mosses: Leucobryum aduncum and Campylopus schmidii. Biodiversitas 21, 2751–2758 (2020).

  25. Asakawa, Y. Biologically active compounds from bryophytes. Pure Appl. Chem. 79, 557–580 (2007).

    Article  Google Scholar 

  26. Bastida, F. et al. Soil microbial diversity–biomass relationships are driven by soil carbon content across global biomes. ISME J. 15, 2081–2091 (2021).

    Article  Google Scholar 

  27. Basile, A., Giordano, S., López-Sáez, J. A. & Cobianchi, R. C. Antibacterial activity of pure flavonoids isolated from mosses. Phytochemistry 52, 1479–1482 (1999).

    Article  Google Scholar 

  28. Rousk, K., Jones, D. L. & DeLuca, T. H. Moss–cyanobacteria associations as biogenic sources of nitrogen in boreal forest ecosystems. Front. Microbiol. (2013).

  29. Commisso, M. et al. Bryo-activities: a review on how bryophytes are contributing to the arsenal of natural bioactive compounds against fungi. Plants (2021).

  30. Carter, D. & Arocena, J. Soil formation under two moss species in sandy materials of central British Columbia (Canada). Geoderma 98, 157–176 (2000).

    Article  Google Scholar 

  31. Glime, J. M. Bryophyte Ecology Vol. 1 (Michigan Tech. Univ. and Int. Assoc. Bryol., 2017).

  32. Herlemann, D. P. R. et al. Transitions in bacterial communities along the 2000 km salinity gradient of the Baltic Sea. ISME J. 5, 1571–1579 (2011).

    Article  Google Scholar 

  33. Ihrmark, K. et al. New primers to amplify the fungal ITS2 region—evaluation by 454-sequencing of artificial and natural communities. FEMS Microb. Ecol. 82, 666–677 (2012).

    Article  Google Scholar 

  34. Callahan, B. J. et al. DADA2: high-resolution sample inference from Illumina amplicon data. Nat. Methods 7, 581–583 (2016).

    Article  Google Scholar 

  35. Tedersoo, L. et al. Regional-scale in-depth analysis of soil fungal diversity reveals strong pH and plant species effects in northern Europe. Front. Microbiol. 11, 1953 (2020).

    Article  Google Scholar 

  36. Bell, C. W. et al. High-throughput fluorometric measurement of potential soil extracellular enzyme activities. J. Vis. Exp. 15, e50961 (2013).

    Google Scholar 

  37. Campbell, C., Chapman, S., Cameron, C., Davidson, M. & Potts, J. A. Rapid microtiter plate method to measure carbon dioxide evolved from carbon substrate amendments so as to determine the physiological profiles of soil microbial communities by using whole soil. Appl. Environ. Microbiol. 69, 3593–3599 (2003).

  38. Frostegard, A. et al. Use and misuse of PLFA measurements in soils. Soil Biol. Biochem. 43, 1621–1625 (2011).

    Article  Google Scholar 

  39. Hu, H., Jung, K., Wang, Q., Saif, L. J., & Vlasova, A. N. Development of a one-step RT-PCR assay for detection of pancoronaviruses (α-, β-, γ-, and δ-coronaviruses) using newly designed degenerate primers for porcine and avian fecal samples. J. Virol. Methods 256, 116–122 (2018).

  40. Schmittgen, T. D. & Livak, K. J. Analyzing real-time PCR data by the comparative C(T) method. Nat. Protoc. 3, 1101–1108 (2008).

    Article  Google Scholar 

  41. Nguyen, N. H. et al. FUNGuild: an open annotation tool for parsing fungal community datasets by ecological guild. Fungal Ecol. 20, 241–248 (2015).

    Article  Google Scholar 

  42. Grace, J. B. Structural Equation Modelling and Natural Systems (Cambridge Univ. Press, 2006).

  43. Le, T. B., Wu, J. & Gong, Y. Vascular plants regulate responses of boreal peatland Sphagnum to climate warming and nitrogen addition. Sci. Total Environ. (2022).

  44. Archer, E. rfPermute: Estimate Permutation p-Values for Random Forest Importance Metrics v.1.5.2 (2016).

  45. Lahouar, A. & Slama, J. B. H. Day-ahead load forecast using random forest and expert input selection. Energy Convers. Manage. 103, 1040–1051 (2015).

    Article  Google Scholar 

  46. Loveland, T. R. et al. An analysis of the IGBP global land-cover characterization process. Photogramm. Eng. Remote Sens. 65, 1021–1032 (1999).

    Google Scholar 

  47. Lembrechts, J. J. et al. Global maps of soil temperature. Glob. Change Biol. 28, 3110–3144 (2021).

  48. Hengl, T. et al. SoilGrids250m: global gridded soil information based on machine learning. PLoS ONE 12, e0169748 (2017).

    Article  Google Scholar 

  49. Piñeiro, G., Perelman, S., Guerschman, J. P. & Paruelo, J. M. How to evaluate models: observed vs. predicted or predicted vs. observed? Ecol. Model. 216, 316–322 (2008).

    Article  Google Scholar 

  50. Brown, S. bivarRasterPlot.R (, accessed 6 June 2022);

  51. Eldridge, D. J. & Delgado-Baquerizo, M. Soil mosses support the delivery of critical ecosystem services globally. Figshare (2023).

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We thank D. Wardle for his insightful comments on an earlier draft and A. Gallardo for his assistance during sample collection. We are grateful to V. Hugonnot, J. G. Segarra-Moragues, F. Müller, S. Stix, I. Charissou, D. Yann, M.-F. Indorf and BRYONET for assistance identifying moss species. We thank S. C. Angorrilla for help with laboratory analyses. The study work associated with this paper was funded by a Large Research Grant from the British Ecological Society (no. LRB17\1019; MUSGONET). D.J.E. is supported by the Hermon Slade Foundation. M.D.-B. was supported by a Ramón y Cajal grant from the Spanish Ministry of Science and Innovation (RYC2018-025483-I), a project from the Spanish Ministry of Science and Innovation for the I + D + i (PID2020-115813RA-I00 funded by MCIN/AEI/10.13039/501100011033a) and a project PAIDI 2020 from the Junta de Andalucía (P20_00879). E.G. is supported by the European Research Council grant agreement 647038 (BIODESERT). M.B. is supported by a Ramón y Cajal grant from Spanish Ministry of Science (RYC2021-031797-I). A.d.l.R is supported by the AEI project PID2019-105469RB-C22. L.W. and Jianyong Wang are supported by the Program for Introducing Talents to Universities (B16011) and the Ministry of Education Innovation Team Development Plan (2013-373). The contributions of T.G. and T.U.N. were supported by the Research Program in Forest Biology, Ecology and Technology (P4-0107) and the research projects J4-3098 and J4-4547 of the Slovenian Research Agency. The contribution of P.B.R. was supported by the NSF Biological Integration Institutes grant DBI-2021898. J. Durán and A. Rodríguez acknowledge support from the FCT (2020.03670.CEECIND and SFRH/BDP/108913/2015, respectively), as well as from the MCTES, FSE, UE and the CFE (UIDB/04004/2021) research unit financed by FCT/MCTES through national funds (PIDDAC).

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Authors and Affiliations



M.D.-B. and D.J.E. developed the original idea of the analyses presented in the paper. M.D.-B. designed the field study and wrote the grant that funded the work. Field data were collected by M.D.-B., D.J.E., M.B., J.L.B.-P., C.P., S.M., S.A., F.A., A.R.B., A.d.l.R., J. Durán., T.G., J.G.I., Y.-R.L., T.P.M., M.A.M.-M. T.U.N., G.F.P.-B., A. Rey, A. Rodriguez, C.S., A.L.T., C.T.-D., P.T., L.W., Jianyong Wang, E.Z., X.Z. and X.-Q.Z. Laboratory analyses were performed by M.D.-B., H.-W.H., J.-Z.H., F.B., J.L.M., L.T., T.S.-S., T.Y., W.S., H.C., S.P. and P.T. Mapping and remote sensing were performed by J.J.G., E.G. and C.A.G. and bioinformatic analyses by B.S., Juntao Wang, H.-W.H. and J.-Z.H. Meta-analytical data were collected by S.L. and G.Z. Statistical analyses were carried out by M.D.-B, J. Ding and M.M.-C. The paper was written by D.J.E. and M.D.-B. and edited by P.B.R., R.O.-H. and J. Ding with contributions from all co-authors.

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Correspondence to David J. Eldridge or Manuel Delgado-Baquerizo.

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Nature Geoscience thanks Bernhard Schmid, Brian Steidinger and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Primary Handling Editor: Xujia Jiang, in collaboration with the Nature Geoscience team.

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Supplementary Figs. 1–19, Tables 1–9 and Appendices 1–5.

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Supplementary Video 1

Video showing a sample of study sites in Google Earth and photographs of their mosses.

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Eldridge, D.J., Guirado, E., Reich, P.B. et al. The global contribution of soil mosses to ecosystem services. Nat. Geosci. 16, 430–438 (2023).

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