An epic global study of moss reveals it is far more vital to Earth’s ecosystems than we knew

Image: Dylan Shaw UnSplash 

May 2, 2023 6.00am AEST in The Conversation

  1. Professor of Dryland Ecology, UNSW Sydney

  2. Ecosystem ecologist, Spanish National Research Council, Consejo Superior de Investigaciones Científicas (CSIC)

Mosses are some of the oldest land plants. They are found all over the world, from lush tropical rainforests to the driest deserts, and even the wind-swept hills of Antarctica.

They are everywhere; growing in cracks along roads and pathways, on the trunks of trees, on rocks and buildings, and importantly, on the soil.

Yet despite this ubiquity, we have a relatively poor understanding of how important they are, particularly the types of moss that thrive on soil.

New global research on soil mosses published today in Nature Geoscience reveals they play critical roles in sustaining life on our planet. Without soil mosses, Earth’s ability to produce healthy soils, provide habitat for microbes and fight pathogens would be greatly diminished.

Soil moss with fruiting bodies (capsules). David Eldridge, Author provided.

A global survey of soil mosses

The results of the new study indicate we have probably underestimated just how important soil mosses are.

Using data from 123 sites across all continents including Antarctica, we show that the soil beneath mosses has more nitrogen, phosphorus and magnesium, and a greater activity of soil enzymes than bare surfaces with no plants.

In fact, mosses affect all major soil functions, increasing carbon sequestration, nutrient cycling and the breakdown of organic matter. These processes are critical for sustaining life on Earth.

Our modelling revealed that soil mosses cover a huge area of the planet, about 9 million square kilometres – equivalent to the area of China. And that’s not counting mosses from boreal forests, which were not included in the study.

The strength of the effect mosses have on soil depends on their growing conditions. They have the strongest effect in natural low productivity environments, such as deserts. They are also more important on sandy and salty soils, and where rainfall is highly variable.

Not unexpectedly, mosses have the strongest effects on soils where vascular plants– those that contain specialised tissues to conduct water and minerals – are sparse.

Read more: Silver moss is a rugged survivor in the city landscape

An intimate connection

Mosses lack the plumbing that allows vascular plants to grow tall and pull water from beneath the soil. This keeps them relatively short, and means they develop an intimate connection with the uppermost soil layers.

Mosses are extremely absorbent and can attract airborne dust particles. Some of these particles are incorporated into the soil below. It is not surprising then that they have such a strong effect on soils.

Our modelling shows that, across the globe, mosses store 6.4 gigatonnes more carbon than soils without plant cover.

Losing just 15% of the global cover of soil mosses would be equivalent to global emissions of carbon dioxide from all land use changes over a year, such as clearing and overgrazing.

A forest floor with rich green moss cover seen in the foreground
Without mosses, the world’s ecosystems wouldn’t thrive nearly as well. Eric Prouzet/Unsplash

Not all mosses are equal

We also found some mosses are more effective at promoting healthy soils than others. Long-lived mosses tended to be associated with more carbon and greater control of soil pathogens.

The ability of mosses to provide ecosystem services and support a diverse community of microbes, fungi and invertebrates was strongest in locations with a high cover of mat- and turf-forming mosses such as Sphagnum, which are widely distributed in boreal forests.

Soils are a huge reservoir of soil pathogens, yet the soil beneath mosses had a lower proportion of plant pathogens. Mosses can help to reduce the pathogen load in soils. This ability may have originated when mosses evolved as land plants.

A special group in the desert

A special type of moss flourishes in deserts. They either live hard (perennial mosses) or die young (annual mosses).

Mosses in the family Pottiaceae are uniquely suited to life under dry and inhospitable conditions. Many have specialised structures that allow them to survive when water is scarce. These include boat-shaped leaves with long hairy tips that help to funnel water into the centre of the plant. Some mosses twist around their stem to reduce the area exposed to the sun and conserve moisture.

Long hair points on the leaves of Campylopus sp. David Eldridge, Author provided

Desert mosses also protect the soil against erosion, influence how much watermoves through the upper layers and even alter the survival chances of plant seedlings.

Other mosses have special moisture-absorbing cells (papillae) that swell up and provide them with a moisture reserve when conditions are dry.

Papillae on the leaf of the moss Crossidium davidai. David Eldridge, Author provided

Our global study showed that mat- and turf-forming mosses such as Sphagnumhad the strongest positive effects on the diversity of microbes, fungi and invertebrates, and on critical services such as nutrient supply. Predictably, longer-lived mosses supported more soil carbon and had greater control of plant pathogens than short-lived mosses.

Protect the mosses

Overall, our work shows mosses influence important soil processes and function in the same way vascular plants do. Their effects may not be as strong, but their total cover means mosses are potentially as significant when summed across the whole globe.

But mosses are under increasing threats globally; disturbance by livestock, overharvesting, land clearing and even changing climates are the greatest threats.

We need a greater acknowledgement of the services that soil mosses provide for all life on this planet. This means greater education about their positive benefits, identifying and mitigating the main threats they face, and including them in routine monitoring programs.

Soil mosses are everywhere, but their future is far from secure. They are likely to play increasingly important roles as vascular plants decline under predicted hotter, drier and more variable global climates.

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 a high cover of mat and turf mosses, in less-productive ecosystems and on sandy and salty 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.


  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 schmidiiBiodiversitas 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 Biol28, 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).

Pledge Your Vote Now
Change language