Analyze Diet
Animals : an open access journal from MDPI2025; 15(6); 862; doi: 10.3390/ani15060862

Semi-Feral Horse Grazing Benefits the Grassland Diversity of Flowering Plants Including a Pollinator-Promoting Indicator Species.

Abstract: European grasslands and their biodiversity are declining rapidly due to land use changes, which highlight the need to develop effective restoration strategies. This study investigates the impact of reintroducing the Swedish national horse breed (the Gotland Russ) on grassland plant diversity and evenness in abandoned agricultural landscapes in Southeast Sweden. Twelve horses were introduced into three 10-13-hectare enclosure replicates (four horses per enclosure) in a three-year (2014-2016) rewilding experiment. Plant species richness, evenness, and diversity were investigated in both grazed and un-grazed conditions. The results indicate that horse grazing significantly increased grassland plant species diversity and richness, with higher Shannon and Simpson's diversity indices in grazed areas. In addition, the abundance of white clover (), a signal species beneficial to pollinators, increased significantly in grazed areas. These findings emphasize the need for integrating large herbivore grazing into ecological restoration practices. Considering the recently enacted EU Nature Restoration Law, which aims to restore 20% of Europe's degraded ecosystems by 2030, this research provides critical insights into scalable restoration methods. The implementation of restoration strategies that include large herbivores may enhance the resilience and biodiversity of European grasslands, thereby aligning with the EU's restoration goals.
Publication Date: 2025-03-17 PubMed ID: 40150391PubMed Central: PMC11939236DOI: 10.3390/ani15060862Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
  • Journal Article

Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

This study explores the potential of using semi-feral horses, specifically the Gotland Russ breed, to restore plant diversity in declining European grasslands. The researchers found that horse grazing in abandoned agricultural landscapes significantly increased the diversity and richness of plant species, including a species favorable to pollinators.

Research Background

  • The research is set against the backdrop of rapidly declining biodiversity in European grasslands due to changes in land use. This situation creates an urgent need to develop practical and effective strategies to restore these ecosystems.
  • The focus of this study is in Southeast Sweden, where abandoned agricultural landscapes are used as experimental sites.

Research Methodology

  • Twelve Gotland Russ horses, a Swedish national breed, were introduced into three enclosures, each ranging from 10 to 13 hectares. Each enclosure was populated by four horses.
  • For three years (2014-2016), the researchers observed the impact of these horses on the diversity, richness, and evenness of plant species, comparing grazed areas with un-grazed ones.

Key Findings

  • The study found that horse grazing resulted in a significant increase in plant species diversity and richness in grazed areas. This outcome was consistent in the various measurements of the Shannon and Simpson’s diversity indices used by the researchers.
  • The researchers also noticed a substantial increase in the abundance of white clover, a species known to be beneficial to pollinators, in grazed areas.

Implications for Ecological Restoration

  • The results highlight the potential benefits of integrating large herbivore grazing into restoration practices. The reintroduction of horses, in this case, helped in enhancing plant biodiversity within the experimental sites.
  • These findings could provide important insights for implementing the recently enacted EU Nature Restoration Law. The law aims to restore 20% of Europe’s degraded ecosystems by 2030, and using large herbivores as part of restoration strategies aligns well with this goal.
  • This research presents a scalable solution that could potentially enhance the resilience and biodiversity of European grasslands, thus, playing an essential part in the fight against biodiversity decline on the continent.

Cite This Article

APA
Thulin CG, Chen Y, Garrido P. (2025). Semi-Feral Horse Grazing Benefits the Grassland Diversity of Flowering Plants Including a Pollinator-Promoting Indicator Species. Animals (Basel), 15(6), 862. https://doi.org/10.3390/ani15060862

Publication

ISSN: 2076-2615
NlmUniqueID: 101635614
Country: Switzerland
Language: English
Volume: 15
Issue: 6
PII: 862

Researcher Affiliations

Thulin, Carl-Gustaf
  • Department of Animal Biosciences, Swedish University of Agricultural Sciences, 750 07 Uppsala, Sweden.
Chen, Yufei
  • Department of Animal Biosciences, Swedish University of Agricultural Sciences, 750 07 Uppsala, Sweden.
Garrido, Pablo
  • Department of Animal Biosciences, Swedish University of Agricultural Sciences, 750 07 Uppsala, Sweden.
  • Natural Capital Foundation (Fundación Patrimonio Natural de Castilla y León), 470 08 Valladolid, Spain.

Grant Funding

  • SWE 0163 / Stiftelsen Vu00e4rldsnaturfonden WWF
  • 160201 / Helge Ax:son Johnsons Stiftelse

Conflict of Interest Statement

The authors declare no conflicts of interest.

References

This article includes 62 references
  1. Malhi Y, Lander T, le Roux E, Stevens N, Macias-Fauria M, Wedding L, Girardin C, Kristensen JÅ, Sandom CJ, Evans TD. The role of large wild animals in climate change mitigation and adaptation.. Curr. Biol. 2022;32:R181–R196.
    doi: 10.1016/j.cub.2022.01.041pubmed: 35231416google scholar: lookup
  2. Smith FA, Elliott Smith RE, Lyons SK, Payne JL, Villaseñor A. The accelerating influence of humans on mammalian macroecological patterns over the late Quaternary.. Quat. Sci. Rev. 2019;211:1–16.
  3. Davoli M, Monsarrat S, Pedersen RØ, Scussolini P, Karger DN, Normand S, Svenning J-C. Megafauna diversity and functional declines in Europe from the Last Interglacial to the present.. Glob. Ecol. Biogeogr. 2024;33:34–47.
    doi: 10.1111/geb.13778google scholar: lookup
  4. Svenning J-C, Lemoine RT, Bergman J, Buitenwerf R, Le Roux E, Lundgren E, Mungi N, Pedersen RØ. The late-Quaternary megafauna extinctions: Patterns, causes, ecological consequences and implications for ecosystem management in the Anthropocene.. Camb. Prism. Extinction. 2024;2:e5.
    doi: 10.1017/ext.2024.4pmc: PMC11895740pubmed: 40078803google scholar: lookup
  5. Pearce EA, Mazier F, Normand S, Fyfe R, Andrieu V, Bakels C, Balwierz Z, Bińka K, Boreham S, Borisova OK. Substantial light woodland and open vegetation characterized the temperate forest biome before Homo sapiens.. Sci. Adv. 2023;9:eadi9135.
    doi: 10.1126/sciadv.adi9135pmc: PMC10637746pubmed: 37948521google scholar: lookup
  6. Barnosky AD, Lindsey EL, Villavicencio NA, Bostelmann E, Hadly EA, Wanket J, Marshall CR. Variable impact of late-Quaternary megafaunal extinction in causing ecological state shifts in North and South America.. Proc. Natl. Acad. Sci. USA. 2016;113:856–861.
    doi: 10.1073/pnas.1505295112pmc: PMC4739530pubmed: 26504219google scholar: lookup
  7. Gill JL. Ecological impacts of the late Quaternary megaherbivore extinctions.. New Phytol. 2014;201:1163–1169.
    doi: 10.1111/nph.12576pubmed: 24649488google scholar: lookup
  8. Cromsigt JPGM, te Beest M, Kerley GIH, Landman M, le Roux E, Smith FA. Trophic rewilding as a climate change mitigation strategy?. Philos. Trans. R. Soc. B Biol. Sci. 2018;373:20170440.
    doi: 10.1098/rstb.2017.0440pmc: PMC6231077pubmed: 30348867google scholar: lookup
  9. Hyvarinen O, te Beest M, le Roux E, Kerley GIH, Buitenwerf R, Druce DJ, Chen J, Rapp L, Fernandes J, Cromsigt JPGM. Megagrazer loss drives complex landscape-scale biophysical cascades.. Environ. Res. Lett. 2025;20:024028.
    doi: 10.1088/1748-9326/ada16egoogle scholar: lookup
  10. Martin PS. Twilight of the Mammoths: Ice Age Extinctions and the Rewilding of America.. University of California Press; Berkley, CA, USA: 2005. pp. 1–269.
  11. Bergman J, Pedersen RØ, Lundgren EJ, Lemoine RT, Monsarrat S, Pearce EA, Schierup MH, Svenning J-C. Worldwide Late Pleistocene and Early Holocene population declines in extant megafauna are associated with Homo sapiens expansion rather than climate change.. Nat. Commun. 2023;14:7679.
    doi: 10.1038/s41467-023-43426-5pmc: PMC10667484pubmed: 37996436google scholar: lookup
  12. Lemoine RT, Buitenwerf R, Svenning J-C. Megafauna extinctions in the late-Quaternary are linked to human range expansion, not climate change.. Anthropocene. 2023;44:100403.
  13. Kavar T, Dovč P. Domestication of the horse: Genetic relationships between domestic and wild horses.. Livest. Sci. 2008;116:1–14.
  14. Librado P, Khan N, Fages A, Kusliy MA, Suchan T, Tonasso-Calvière L, Schiavinato S, Alioglu D, Fromentier A, Perdereau A. The origins and spread of domestic horses from the Western Eurasian steppes.. Nature. 2021;598:634–640.
    doi: 10.1038/s41586-021-04018-9pmc: PMC8550961pubmed: 34671162google scholar: lookup
  15. Harris PA. Developments in equine nutrition: Comparing the beginning and end of this century.. J. Nutr. 1998;128:S2698–S2703.
    doi: 10.1093/jn/128.12.2698Spubmed: 9868245google scholar: lookup
  16. Bignal EM, McCracken DI. Low-intensity farming systems in the conservation of the countryside.. J. Appl. Ecol. 1996 33:413–424.
    doi: 10.2307/2404973google scholar: lookup
  17. Eriksson O, Cousins SAO, Bruun HH. Land-use history and fragmentation of traditionally managed grasslands in Scandinavia.. J. Veg. Sci. 2002;13:743–748.
  18. Garrido P, Edenius L, Mikusiński G, Skarin A, Jansson A, Thulin C-G. Experimental rewilding may restore abandoned wood-pastures if policy allows.. Ambio. 2021;50:101–112.
    doi: 10.1007/s13280-020-01320-0pmc: PMC7708577pubmed: 32152907google scholar: lookup
  19. Mutillod C, Buisson E, Tatin L, Mahy G, Dufrêne M, Mesléard F, Dutoit T. Managed as wild, horses influence grassland vegetation differently than domestic herds.. Biol. Conserv. 2024;290:110469.
  20. Valdés-Correcher E, Sitters J, Wassen M, Brion N, Olde Venterink H. Herbivore dung quality affects plant community diversity.. Sci. Rep. 2019;9:5675.
    doi: 10.1038/s41598-019-42249-zpmc: PMC6450897pubmed: 30952928google scholar: lookup
  21. Campbell JE, Gibson DJ. The effect of seeds of exotic species transported via horse dung on vegetation along trail corridors.. Plant Ecol. 2001;157:23–35.
    doi: 10.1023/A:1013751615636google scholar: lookup
  22. Öckinger E, Eriksson AK, Smith HG. Effects of grassland abandonment, restoration and management on butterflies and vascular plants.. Biol. Conserv. 2006;133:291–300.
  23. Garrido P, Mårell A, Öckinger E, Skarin A, Jansson A, Thulin C-G. Experimental rewilding enhances grassland functional composition and pollinator habitat use.. J. Appl. Ecol. 2019;56:946–955.
    doi: 10.1111/1365-2664.13338google scholar: lookup
  24. Garrido P, Thulin C-G, Negro JJ. EU nature restoration law fails to recognize missing large herbivore functions.. Biol. Conserv. 2025;303:111026.
  25. Svenning J-C, Pedersen PBM, Donlan CJ, Ejrnæs R, Faurby S, Galetti M, Hansen DM, Sandel B, Sandom CJ, Terborgh JW. Science for a wilder Anthropocene: Synthesis and future directions for trophic rewilding research.. Proc. Natl. Acad. Sci. USA. 2016;113:898–906.
    doi: 10.1073/pnas.1502556112pmc: PMC4743824pubmed: 26504218google scholar: lookup
  26. Ringmark S, Skarin A, Jansson A. Impact of year-round grazing by horses on pasture nutrient dynamics and the correlation with pasture nutrient content and fecal nutrient composition.. Animals. 2019;9:500.
    doi: 10.3390/ani9080500pmc: PMC6720502pubmed: 31362460google scholar: lookup
  27. Chen Y. Impact of Horses on Year Around Grazing Without Supplementary Feeding on Pastoral Herbaceous Plants.. Master’s Thesis. Swedish University of Agricultural Sciences; Uppsala, Sweden: 2024.
  28. Tydén E, Jansson A, Ringmark S. Parasites in Horses Kept in A 2.5 Year-Round Grazing System in Nordic Conditions without Supplementary Feeding.. Animals. 2019;9:1156.
    doi: 10.3390/ani9121156pmc: PMC6940839pubmed: 31861066google scholar: lookup
  29. Oksanen J, Simpson GL, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, Solymos P, Stevens MHH, Szoecs E. Vegan: Community Ecology Package (2.6-4). [(accessed on 12 February 2025)].
  30. Bates D, Mächler M, Bolker B, Walker S. Fitting linear mixed-effects models using lme4.. J. Stat. Softw. 2015;67:1–48.
    doi: 10.18637/jss.v067.i01google scholar: lookup
  31. Lawton JH, Gaston KJ. Indicator species.. Encycl. Biodivers. 2021;3:437–450.
  32. Egan LM, Hofmann RW, Ghamkhar K, Hoyos-Villegas V. Prospects for Trifolium improvement through germplasm characterisation and pre-breeding in New Zealand and beyond.. Front. Plant Sci. 2021;12:653191.
    doi: 10.3389/fpls.2021.653191pmc: PMC8242581pubmed: 34220882google scholar: lookup
  33. Hyslop MG, Kemp PD, Hodgson J. Vegetatively reproductive red clovers (Trifolium pratense L.): An overview.. Proc. N. Z. Grassl. Assoc. 1999;61:121–126.
  34. Sawicka B, Krochmal-Marczak B, Sawicki J, Skiba D, Pszczółkowski P, Barbaś P, Vambol V, Messaoudi M, Farhan AK. White clover (Trifolium repens L.) cultivation as a means of soil regeneration and pursuit of a sustainable food system model.. Land. 2023;12:838.
    doi: 10.3390/land12040838google scholar: lookup
  35. Beye H, Taube F, Lange K, Hasler M, Kluß C, Loges R, Diekötter T. Species-enriched grass-clover mixtures can promote bumblebee abundance compared with intensively managed conventional pastures.. Agronomy. 2022;12:1080.
    doi: 10.3390/agronomy12051080google scholar: lookup
  36. R Core Team. R: The R Project for Statistical Computing.. [(accessed on 12 February 2025)].
  37. Marion B, Bonis A, Bouzillé J-B. How much does grazing-induced heterogeneity impact plant diversity in wet grasslands?. Écoscience. 2010;17:229–239.
    doi: 10.2980/17-3-3315google scholar: lookup
  38. Li W, Liu C, Wang W, Zhou H, Xue Y, Xu J, Xue P, Yan H. Effects of different grazing disturbances on the plant diversity and ecological functions of alpine grassland ecosystem on the Qinghai-Tibetan Plateau.. Front. Plant Sci. 2021;12:765070.
    doi: 10.3389/fpls.2021.765070pmc: PMC8710682pubmed: 34966399google scholar: lookup
  39. Stewart GB, Pullin AS. The relative importance of grazing stock type and grazing intensity for conservation of mesotrophic ‘old meadow’ pasture.. J. Nat. Conserv. 2008;16:175–185.
    doi: 10.1016/j.jnc.2008.09.005google scholar: lookup
  40. Škornik S, Vidrih M, Kaligarič M. The effect of grazing pressure on species richness, composition and productivity in North Adriatic Karst pastures.. Plant Biosyst.-Int. J. Deal. All Asp. Plant Biol. 2010;144:355–364.
    doi: 10.1080/11263501003750250google scholar: lookup
  41. Bonavent C, Olsen K, Ejrnæs R, Fløjgaard C, Hansen MDD, Normand S, Svenning J-C, Bruun HH. Grazing by semi-feral cattle and horses supports plant species richness and uniqueness in grasslands.. Appl. Veg. Sci. 2023;26:e12718.
    doi: 10.1111/avsc.12718google scholar: lookup
  42. Köhler M, Hiller G, Tischew S. Year-round horse grazing supports typical vascular plant species, orchids and rare bird communities in a dry calcareous grassland.. Agric. Ecosyst. Environ. 2016;234:48–57.
  43. Beck JJ, Hernández DL, Pasari JR, Zavaleta ES. Grazing maintains native plant diversity and promotes community stability in an annual grassland.. Ecol. Appl. 2015;25:1259–1270.
    doi: 10.1890/14-1093.1pubmed: 26485954google scholar: lookup
  44. Huisman J, Olff H. Competition and facilitation in multispecies plant-herbivore systems of productive environments.. Ecol. Lett. 1998;1:25–29.
  45. Jutila HM, Grace JB. Effects of disturbance on germination and seedling establishment in a coastal prairie grassland: A test of the competitive release hypothesis.. J. Ecol. 2002;90:291–302.
  46. Lundgren EJ, Schowanek SD, Rowan J, Middleton O, Pedersen RØ, Wallach AD, Ramp D, Davis M, Sandom CJ, Svenning J-C. Functional traits of the world’s late Quaternary large-bodied avian and mammalian herbivores.. Sci. Data. 2021;8:17.
    doi: 10.1038/s41597-020-00788-5pmc: PMC7817692pubmed: 33473149google scholar: lookup
  47. Garrido P, Naumov V, Söderquist L, Jansson A, Thulin C-G. Effects of experimental rewilding on butterflies, bumblebees and grasshoppers.. J. Insect Conserv. 2022;26:763–771.
  48. van Klink R, de Vries MFW. Risks and opportunities of trophic rewilding for arthropod communities.. Philos. Trans. R. Soc. B Biol. Sci. 2018;373:20170441.
    doi: 10.1098/rstb.2017.0441pmc: PMC6231068pubmed: 30348868google scholar: lookup
  49. van Klink R, van der Plas F, Van Noordwijk C, WallisDeVries MFW, Olff H. Effects of large herbivores on grassland arthropod diversity.. Biol. Rev. 2015;90:347–366.
    doi: 10.1111/brv.12113pmc: PMC4402009pubmed: 24837856google scholar: lookup
  50. Huang J, Garrido P, Thulin C-G. Experimental rewilding effects on grasslands detected through aerial photographs.. Oecologia. (submitted).
  51. Tälle M, Ranius T, Öckinger E. The usefulness of surrogates in biodiversity conservation: A synthesis.. Biol. Conserv. 2023;288:110384.
  52. Liu J, Feng C, Wang D, Wang L, Wilsey BJ, Zhong Z. Impacts of grazing by different large herbivores in grassland depend on plant species diversity.. J. Appl. Ecol. 2015;52:1053–1062.
    doi: 10.1111/1365-2664.12456google scholar: lookup
  53. Kohyani P, Bossuyt B, Bonte D, Hoffmann M. Grazing impact on plant spatial distribution and community composition.. Plant Ecol. Evol. 2011;144:19–28.
    doi: 10.5091/plecevo.2011.429google scholar: lookup
  54. Li Y, Dong S, Gao Q, Fan C, Fayiah M, Ganjurjav H, Hu G, Wang X, Yan Y, Gao X. Grazing changed plant community composition and reduced stochasticity of soil microbial community assembly of alpine grasslands on the Qinghai-Tibetan Plateau.. Front. Plant Sci. 2022;13:864085.
    doi: 10.3389/fpls.2022.864085pmc: PMC9168915pubmed: 35677251google scholar: lookup
  55. Pywell RF, Warman EA, Carvell C, Sparks TH, Dicks LV, Bennett D, Wright A, Critchley CNR, Sherwood A. Providing foraging resources for bumblebees in intensively farmed landscapes.. Biol. Conserv. 2005;121:479–494.
  56. CBD Sweden. Fifth National Report to the Convention on Biological Diversity-Sweden.. 2014.
  57. Cousins SAO, Auffret AG, Lindgren J, Tränk L. Regional-scale land-cover change during the 20th century and its consequences for biodiversity.. Ambio. 2015;44:17–27.
    doi: 10.1007/s13280-014-0585-9pmc: PMC4288995pubmed: 25576277google scholar: lookup
  58. Sandström J, Carlberg T, Sundberg S, Bjelke U. Tillstånd och Trender för Arter och Deras Livsmiljöer–Rödlistade Arter i Sverige 2015.. ArtDatabanken SLU; Uppsala, Sweden: 2015.
  59. Lasanta T, Nadal-Romero E, Arnáez J. Managing abandoned farmland to control the impact of re-vegetation on the environment. The state of the art in Europe.. Environ. Sci. Policy. 2015;52:99–109.
  60. The Swedish Board of Agriculture. Horses and Horse Establishments in 2016.. Statistiska Meddelanden JO 24 SM 1701 2016, pp. 1–23.
  61. . Regulation (EU) 2024/1991 of the European Parliament and of the Council of 24 June 2024 on Nature Restoration and Amending Regulation (EU) 2022/869.. .
  62. Schmitz OJ, Sylvén M, Atwood TB, Bakker ES, Berzaghi F, Brodie JF, Cromsigt JPGM, Davies AB, Leroux SJ, Schepers FJ. Trophic rewilding can expand natural climate solutions.. Nat. Clim. Change. 2023;13:324–333.

Citations

This article has been cited 0 times.