FREE SHIPPING over $40
OVER 10000 FIVE-STAR REVIEWS
5% OFF ALL SUBSCRIPTIONS
100% HAPPINESS GUARANTEE
Analyze Diet
0
Ecological applications : a publication of the Ecological Society of America2026; 36(1); e70170; doi: 10.1002/eap.70170

Space-use by feral cattle and horses shapes vegetation structure in a trophic rewilding area.

Abstract: Feral cattle (Bos taurus) and horses (Equus ferus caballus) are commonly introduced to European rewilding areas to halt vegetation succession and to conserve light-demanding species. Yet, we still do not understand how the habitat preference of animals shapes vegetation structure at the landscape scale. Here, we used spatial preference modeling to understand drivers of space-use based on GPS-collared horses and cattle in a 120-ha rewilding area in Denmark. Using a time series of a satellite-based vegetation productivity index, we tested the ability of animal space-use to explain changes in vegetation, as well as the trend of its spatial variability at the reserve scale, as a measure of landscape-scale vegetation heterogeneity. We expected that animal space-use would be driven mainly by topography and vegetation characteristics and that highly used areas with open vegetation would remain open. We, indeed, found that vegetation density and landscape connectivity were good predictors of space-use preference for both cattle and horses. Additionally, both cattle and horses were strongly attracted to an artificial shelter located inside the reserve, warranting consideration of the use and placement of artificial infrastructure. Space-use diverged during periods of resource scarcity emphasizing the value of introducing a variety of herbivore functional types for optimizing structural ecosystem heterogeneity. As expected, we found that cattle and horses slow down vegetation succession in highly used areas, as shown by the negative correlation between changes in growing season productivity and intensively used areas dominated by short herbaceous and shrubby vegetation. We could also show that the highly used areas showed the largest reductions and the fastest recovery in vegetation greenness following the pan-European drought in 2018. A ~2/3 reduction in herbivore population size subsequent to the drought was followed by a general greening of the landscape, but with no clear relationship with space-use intensity. Our study supports that trophic rewilding with year-round grazing can limit vegetation densification at the landscape scale under near-natural conditions. This is pertinent in the face of accelerating succession toward increasingly dark and tree-dominated vegetation in temperate Europe's natural areas, and the associated biodiversity loss.
© 2026 The Author(s). Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.
Get Full Text
Publication Date: 2026-02-04 PubMed ID: 41636694PubMed Central: PMC12871458DOI: 10.1002/eap.70170Google 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.
LinkedInCopy
  • 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.

Overview

  • This research examined how the spatial behavior of feral cattle and horses influences vegetation structure in a rewilding area in Denmark.
  • The study aimed to understand animal habitat preferences, their impact on vegetation succession, and landscape-scale vegetation heterogeneity through GPS data and satellite vegetation indices.

Study Context and Objectives

  • Feral cattle (Bos taurus) and horses (Equus ferus caballus) are introduced into European rewilding areas to prevent vegetation succession, helping preserve light-demanding plant species.
  • Despite their use, little was known about how animal habitat preferences shape vegetation patterns over large scales.
  • The study focused on a 120-hectare rewilding reserve in Denmark, leveraging GPS collars on cattle and horses to track space-use.
  • The objectives included:
    • Identifying drivers behind animal spatial preferences (e.g., topography, vegetation features)
    • Determining how animal space-use relates to changes in vegetation productivity and spatial heterogeneity
    • Understanding the impact of grazing on slowing vegetation succession and promoting ecosystem diversity

Methods

  • GPS-collared feral horses and cattle were monitored to gather detailed data on their spatial movements within the reserve.
  • Time-series data from satellite-based vegetation productivity indices (likely NDVI or similar) were used to track vegetation greenness and productivity over time.
  • Statistical modeling linked animal space-use intensity to environmental factors:
    • Topography and vegetation density/connectivity were assessed as predictors of where animals preferred to be.
    • Animal attraction to human-built features, such as an artificial shelter, was also evaluated.
  • Changes in vegetation productivity over growing seasons were analyzed to determine the effects of grazing on succession and heterogeneity.
  • A significant drought in 2018 provided a natural experiment to assess vegetation responses to stress and subsequent recovery under grazing pressure.

Key Findings

  • Vegetation density and landscape connectivity were strong predictors of cattle and horse space-use, implying animals prefer certain vegetation structures.
  • Both species showed a strong attraction to an artificial shelter within the reserve; this highlights the influence of infrastructure on animal movement and habitat use.
  • Differences in space-use emerged during resource scarcity periods, supporting the idea that introducing various herbivore types can enhance ecosystem structural diversity.
  • Heavily grazed areas dominated by short herbaceous and shrubby vegetation experienced slower vegetation succession, indicated by a negative correlation between grazing intensity and increased vegetation productivity.
  • Following the 2018 pan-European drought, areas with high herbivore use showed the largest declines in vegetation greenness but also the fastest recovery, indicating resilience supported by grazing.
  • A substantial reduction (~two-thirds) in herbivore populations after the drought led to overall landscape greening, but no clear link was found between this greening and space-use intensity.

Implications and Conclusions

  • Trophic rewilding with year-round feral grazing can effectively slow vegetation densification and succession across landscapes under near-natural conditions.
  • This addresses a critical conservation challenge: temperate European natural areas are experiencing increased tree domination and denser vegetation, threatening biodiversity.
  • Introducing and managing diverse herbivore species in rewilding areas is important for maintaining heterogeneous vegetation structures that support biodiversity.
  • The strong attraction of animals to human-made features suggests careful planning of infrastructure is needed to avoid unintended effects on grazing patterns.
  • Ultimately, this research supports the role of feral herbivores in shaping vegetation dynamics and promoting landscape-level heterogeneity in rewilding efforts.

Cite This Article

APA
Kristensen JÅ, Buitenwerf R, Berti E, Hansen OLP, Schowanek SD, Ejrnæs R, Hansen MDD, Olsen K, Normand S, Svenning JC. (2026). Space-use by feral cattle and horses shapes vegetation structure in a trophic rewilding area. Ecol Appl, 36(1), e70170. https://doi.org/10.1002/eap.70170

Publication

Ecological applications : a publication of the Ecological Society of America
ISSN: 1939-5582
NlmUniqueID: 9889808
Country: United States
Language: English
Volume: 36
Issue: 1
Pages: e70170
PII: e70170

Researcher Affiliations

Kristensen, Jeppe Å
  • Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Center for Sustainable Landscapes Under Global Change (SustainScapes), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Section for Ecoinformatics and Biodiversity, Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Leverhulme Centre for Nature Recovery, School of Geography and the Environment, University of Oxford, Oxford, UK.
Buitenwerf, Robert
  • Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Section for Ecoinformatics and Biodiversity, Department of Biology, Aarhus University, Aarhus C, Denmark.
Berti, Emilio
  • German Centre for Integrative Biodiversity Research (iDiv), Leipzig, Germany.
  • Institute of Biodiversity, Friedrich-Schiller-University Jena, Jena, Germany.
Hansen, Oskar L P
  • Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Section for Ecoinformatics and Biodiversity, Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Department of Research and Collections, Natural History Museum Aarhus, Aarhus C, Denmark.
Schowanek, Simon D
  • Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Section for Ecoinformatics and Biodiversity, Department of Biology, Aarhus University, Aarhus C, Denmark.
Ejrnæs, Rasmus
  • Department of Ecoscience, Aarhus University, Aarhus C, Denmark.
Hansen, Morten D D
  • Department of Research and Collections, Natural History Museum Aarhus, Aarhus C, Denmark.
Olsen, Kent
  • Department of Research and Collections, Natural History Museum Aarhus, Aarhus C, Denmark.
Normand, Signe
  • Center for Sustainable Landscapes Under Global Change (SustainScapes), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Section for Ecoinformatics and Biodiversity, Department of Biology, Aarhus University, Aarhus C, Denmark.
Svenning, Jens-Christian
  • Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Center for Sustainable Landscapes Under Global Change (SustainScapes), Department of Biology, Aarhus University, Aarhus C, Denmark.
  • Section for Ecoinformatics and Biodiversity, Department of Biology, Aarhus University, Aarhus C, Denmark.

MeSH Terms

  • Animals
  • Cattle / physiology
  • Horses / physiology
  • Conservation of Natural Resources
  • Ecosystem
  • Denmark
  • Plants / classification

Grant Funding

  • 2017-N-10 / 15. Juni Fonden
  • NNF20OC0059595 / Novo Nordisk Fonden
  • FORM.2016-0025 / Danish Agency for Culture and Palaces
  • 6171-00034B / Innovationsfonden
  • DNRF173 / Danmarks Grundforskningsfond
  • CF16-0005 / Carlsbergfondet
  • CF20-0238 / Carlsbergfondet
  • CF23-0641 / Carlsbergfondet
  • 16549 / Villum Fonden
  • 0135-00225B / Danmarks Frie Forskningsfond

Conflict of Interest Statement

The authors declare no conflicts of interest.

References

This article includes 78 references
  1. Allred BW, Fuhlendorf SD, Hamilton RG. The Role of Herbivores in Great Plains Conservation: Comparative Ecology of Bison and Cattle. Ecosphere 2: 1–17.
  2. Assmann JJ, Moeslund JE, Treier UA, Normand S. EcoDes‐DK15: High‐Resolution Ecological Descriptors of Vegetation and Terrain Derived from Denmark's National Airborne Laser Scanning Data Set. Earth System Science Data 14: 823–844.
  3. Auffret AG, Svenning J‐C. Climate Warming Has Compounded Plant Responses to Habitat Conversion in Northern Europe. Nature Communications 13: 7818.
    pmc: PMC9763501pubmed: 36535960
  4. Bergin MD, Pedersen RØ, Jensen M, Svenning J‐C. Mapping Rewilding Potential – A Systematic Approach to Prioritise Areas for Rewilding in Human‐Dominated Regions. Journal for Nature Conservation 77: 126536.
  5. Berti E, Davoli M, Buitenwerf R, Dyer A, Hansen OLP, Hirt M, Svenning JC, Terlau JF, Brose U, Vollrath F. The R Package Enerscape: A General Energy Landscape Framework for Terrestrial Movement Ecology. Methods in Ecology and Evolution 13: 60–67.
  6. Berti E, Kristensen JA, Svenning J‐C. Space‐Use by Feral Cattle and Horses Shapes Vegetation Structure in a Trophic Rewilding Area. Zenodo .
    doi: 10.5281/zenodo.17376061google scholar: lookup
  7. Berti E, Rosenbaum B, Vollrath F. Energy Landscapes Direct the Movement Preferences of Elephants. Journal of Animal Ecology 94: 908–918.
    pmc: PMC12056349pubmed: 40133749
  8. Bocherens H. The Rise of the Anthroposphere since 50,000 Years: An Ecological Replacement of Megaherbivores by Humans in Terrestrial Ecosystems?. Frontiers in Ecology and Evolution 6: 3.
  9. Bonavent C, Olsen K, Ejrnæs R, Fløjgaard C, Hansen MD, Normand S, Svenning J, Bruun HH. Grazing by Semi‐Feral Cattle and Horses Supports Plant Species Richness and Uniqueness in Grasslands. Applied Vegetation Science 26: e12718.
  10. Borowik T, Pettorelli N, Sönnichsen L, Jędrzejewska B. Normalized Difference Vegetation Index (NDVI) as a Predictor of Forage Availability for Ungulates in Forest and Field Habitats. European Journal of Wildlife Research 59: 675–682.
  11. Bråthen KA, Pugnaire FI, Bardgett RD. The Paradox of Forbs in Grasslands and the Legacy of the Mammoth Steppe. Frontiers in Ecology and the Environment 19: 584–592.
  12. Bühne HS, Ross B, Sandom CJ, Pettorelli N. Monitoring Rewilding from Space: The Knepp Estate as a Case Study. Journal of Environmental Management 312: 114867.
    pubmed: 35378468
  13. Buitenwerf R, Sandel B, Normand S, Mimet A, Svenning J‐C. Land Surface Greening Suggests Vigorous Woody Regrowth throughout European Semi‐Natural Vegetation. Global Change Biology 24: 5789–5801.
    pubmed: 30238566
  14. Clauss M, Frey R, Kiefer B, Lechner‐Doll M, Loehlein W, Polster C, Rössner GE, Streich WJ. The Maximum Attainable Body Size of Herbivorous Mammals: Morphophysiological Constraints on Foregut, and Adaptations of Hindgut Fermenters. Oecologia 136: 14–27.
    pubmed: 12712314
  15. Cromsigt JPGM, Kemp YJM, Rodriguez E, Kivit H. Rewilding Europe's Large Grazer Community: How Functionally Diverse Are the Diets of European Bison, Cattle, and Horses?. Restoration Ecology 26: 891–899.
  16. Czyżewski S, Svenning J-C. Temperate Forest Plants Are Associated with Heterogeneous Semi‐Open Canopy Conditions Shaped by Large Herbivores. Nature Plants 11: 985–1000.
    pubmed: 40229585
  17. Davoli M, Monsarrat S, Pedersen R Ø, Scussolini P, Karger D N, Normand S, Svenning J-C. Megafauna Diversity and Functional Declines in Europe from the Last Interglacial to the Present. Global Ecology and Biogeography 33: 34–47.
  18. Dvorský M, Mudrák O, Doležal J, Jirků M. Reintroduction of Large Herbivores Restored Plant Species Richness in Abandoned Dry Temperate Grassland. Plant Ecology 223: 525–535.
  19. Ejrnæs D D, Olivier B, Bakker E S, Cornelissen P, Ejrnæs R, Smit C, Svenning J-C. Vegetation Dynamics Following Three Decades of Trophic Rewilding in the Mesic Grasslands of Oostvaardersplassen. Applied Vegetation Science 27: e12805.
  20. El Garroussi S, Di Giuseppe F, Barnard C, Wetterhall F. Europe Faces up to Tenfold Increase in Extreme Fires in a Warming Climate. npj Climate and Atmospheric Science 7: 30.
  21. Eskelinen A, Harpole W S, Jessen M-T, Virtanen R, Hautier Y. Light Competition Drives Herbivore and Nutrient Effects on Plant Diversity. Nature 611: 301–305.
    pmc: PMC9646529pubmed: 36323777
  22. Estrada-Carmona N, Sánchez A C, Remans R, Jones S K. Complex Agricultural Landscapes Host More Biodiversity than Simple Ones: A Global Meta‐Analysis. Proceedings of the National Academy of Sciences of the United States of America 119: e2203385119.
    pmc: PMC9499564pubmed: 36095174
  23. Fløjgaard C, Pedersen P B M, Sandom C J, Svenning J-C, Ejrnæs R. Exploring a Natural Baseline for Large‐Herbivore Biomass in Ecological Restoration. Journal of Applied Ecology 59: 18–24.
  24. Gashchak S, Paskevych S. Przewalski's Horse () in the Chornobyl Exclusion Zone after 20 Years of Introduction. Theriologia Ukrainica 18: 80–100.
  25. Gilhaus K, Stelzner F, Hölzel N. Cattle Foraging Habits Shape Vegetation Patterns of Alluvial Year‐Round Grazing Systems. Plant Ecology 215: 169–179.
  26. . Sable Island National Park Reserve. .
  27. Hagstrup M, Bruhn D, Olsen K, Lukassen M, Pertoldi C. Molecular Study of Dietary Diversity of the Exmoor‐Ponies (). Genetics and Biodiversity Journal 4: 53–70.
  28. Henning K, Lorenz A, von Oheimb G, Härdtle W, Tischew S. Year‐Round Cattle and Horse Grazing Supports the Restoration of Abandoned, Dry Sandy Grassland and Heathland Communities by Supressing Calamagrostis Epigejos and Enhancing Species Richness. Journal for Nature Conservation 40: 120–130.
  29. Hopcraft J G C, Olff H, Sinclair A R E. Herbivores, Resources and Risks: Alternating Regulation along Primary Environmental Gradients in Savannas. Trends in Ecology & Evolution 25: 119–128.
    pubmed: 19767121
  30. Janis C. The Evolutionary Strategy of the Equidae and the Origins of Rumen and Cecal Digestion. Evolution 30: 757–774.
    pubmed: 28563331
  31. Kleppel GS, Frank DA. Structure and Functioning of Wild and Agricultural Grazing Ecosystems: A Comparative Review. Frontiers in Sustainable Food Systems 6: 945514.
  32. Klich D. Selective Bark Stripping of Various Tree Species by Polish Horses in Relation to Bark Detachability. Forest Ecology and Management 384: 65–71.
  33. Klich D, Stereńczak K, Lisiewicz M, Sobczuk M, Nieszała A, Olech W. An Assessment of the Habitat Preferences of European Bison with Airborne Laser Scanning Data in Forest Ecosystem. Scientific Reports 13: 17987.
    pmc: PMC10589212pubmed: 37864104
  34. 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. Agriculture, Ecosystems & Environment 234: 48–57.
  35. Kovács L. Feature Selection Algorithms in Generalized Additive Models under Concurvity. Computational Statistics 39: 1–33.
  36. Kowalczyk R, Kamiński T, Borowik T. Do Large Herbivores Maintain Open Habitats in Temperate Forests?. Forest Ecology and Management 494: 119310.
  37. Kristensen JA. Space‐Use by Feral Cattle and Horses Shapes Vegetation Structure in a Trophic Rewilding area_NDVIchange. Ecological Applications .
    doi: 10.5281/zenodo.17896843google scholar: lookup
  38. Kullberg Y, Bergström R. Winter Browsing by Large Herbivores on Planted Deciduous Seedlings in Southern Sweden. Scandinavian Journal of Forest Research 16: 371–378.
  39. Liu Y, Zhang Y, Peñuelas J, Kannenberg SA, Gong H, Yuan W, Wu C, Zhou S, Piao S. Drought Legacies Delay Spring Green‐Up in Northern Ecosystems. Nature Climate Change 15: 444–451.
  40. Lundgren EJ, Ramp D, Middleton OS, Wooster EIF, Kusch E, Balisi M, Ripple WJ. A Novel Trophic Cascade between Cougars and Feral Donkeys Shapes Desert Wetlands. Journal of Animal Ecology 91: 2348–2357.
    pmc: PMC10087508pubmed: 35871769
  41. Mas‐Carrió E, Churski M, Kuijper D, Fumagalli L. Niche Overlap across Landscape Variability in Summer between Two Large Herbivores Using eDNA Metabarcoding. PLoS One 19: 1–16.
    pmc: PMC10863879pubmed: 38349911
  42. Mata JC, Svenning J‐C, Chequín RN, Davalos M, Salas R, Vucko A, Buitenwerf R. Capybaras () Generate Grazing Lawns and Maintain Plant Diversity in Neotropical Savanna. Applied Vegetation Science 28: e70013.
  43. Moeslund JE, Arge L, Bøcher PK, Dalgaard T, Odgaard MV, Nygaard B, Svenning J‐C. Topographically Controlled Soil Moisture Is the Primary Driver of Local Vegetation Patterns across a Lowland Region. Ecosphere 4: art91.
  44. Moeslund JE, Arge L, Bøcher PK, Dalgaard T, Svenning J‐C. Topography as a Driver of Local Terrestrial Vascular Plant Diversity Patterns. Nordic Journal of Botany 31: 129–144.
  45. 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. Biological Conservation 290: 110469.
  46. Nelson RA, Sullivan LL, Hersch‐Green EI, Seabloom EW, Borer ET, Tognetti PM, Adler PB. Forb Diversity Globally Is Harmed by Nutrient Enrichment but Can be Rescued by Large Mammalian Herbivory. Communications Biology 8: 444.
    pmc: PMC11910660pubmed: 40089613
  47. Nicodemo MLF, Porfírio‐da‐Silva V. Bark Stripping by Cattle in Silvopastoral Systems. Agroforestry Systems 93: 305–315.
  48. Nieszała A, Klich D, Perzanowski K, Januszczak M, Wołoszyn‐Gałęza A, Olech W. Debarking Intensity of European Bison in the Bieszczady Mountains in Relation to Forest Habitat Features. Forest Ecology and Management 508: 120057.
  49. Olff H, Vera FWM, Bokdam J, Bakker ES, Gleichman JM, de Maeyer K, Smit R. Shifting Mosaics in Grazed Woodlands Driven by the Alternation of Plant Facilitation and Competition. Plant Biology 1: 127–137.
  50. Pausas JG, Bond WJ. Feedbacks in Ecology and Evolution. Trends in Ecology & Evolution 37: 637–644.
    pubmed: 35466019
  51. Pearce EA, Mazier F, Fyfe R, Davison CW, Normand S, Serge M‐A, Scussolini P, Svenning J‐C. Higher Abundance of Disturbance‐Favoured Trees and Shrubs in European Temperate Woodlands Prior to the Late‐Quaternary Extinction of Megafauna. Journal of Ecology 112: 2813–2827.
  52. Pearce EA, Mazier F, Normand S, Fyfe R, Andrieu V, Bakels C, Balwierz Z. Substantial Light Woodland and Open Vegetation Characterized the Temperate Forest Biome before. Science Advances 9: eadi9135.
    pmc: PMC10637746pubmed: 37948521
  53. Popp A, Scheibe KM. The Ecological Influence of Large Herbivores – Behavior and Habitat Utilization of Cattle and Horses. Applied Ecology and Environmental Research 12: 681–693.
  54. Pratt RM, Putman RJ, Ekins JR, Edwards PJ. Use of Habitat by Free‐Ranging Cattle and Ponies in the New Forest, Southern England. The Journal of Applied Ecology 23: 539–557.
  55. Putman RJ, Pratt RM, Ekins JR, Edwards PJ. Food and Feeding Behaviour of Cattle and Ponies in the New Forest, Hampshire. The Journal of Applied Ecology 24: 369–380.
  56. R Core Team. R: A Language and Environment for Statistical Computing. Vienna: R Foundation for Statistical Computing.
  57. Ratajczak Z, Collins SL, Blair JM, Nippert JB. Reintroducing Bison Results in Long‐Running and Resilient Increases in Grassland Diversity. Proceedings of the National Academy of Sciences of the United States of America 119: e2210433119.
    pmc: PMC9457053pubmed: 36037376
  58. Rech BJ, Buitenwerf R, Ruggiero R, Trepel J, Waltert M, Svenning J‐C. What Moves Large Grazers? Habitat Preferences and Complementing Niches of Large Herbivores in a Danish Trophic Rewilding Area. Environmental Management 75: 1665–1679.
    pmc: PMC12228648pubmed: 40278902
  59. Sandom CJ, Ejrnaes R, Hansen MDD, Svenning JC. High Herbivore Density Associated with Vegetation Diversity in Interglacial Ecosystems. Proceedings of the National Academy of Sciences of the United States of America 111: 4162–4167.
    pmc: PMC3964052pubmed: 24591633
  60. Scasta JD, Beck JL, Angwin CJ. Meta‐Analysis of Diet Composition and Potential Conflict of Wild Horses with Livestock and Wild Ungulates on Western Rangelands of North America. Rangeland Ecology & Management 69: 310–318.
  61. Smagol V, Khoyetskyy P, Yarysh V, Smagol V, Maievskyi K, Plumb G. Habitat Characteristics of European Bison () in Ukraine. European Journal of Wildlife Research 68: 29.
  62. Søndergaard SA, Ejrnæs R, Svenning J-C, Fløjgaard C. From Grasslands to Forblands: Year‐Round Grazing as a Driver of Plant Diversity. Journal of Applied Ecology 62: 1104–1113.
  63. Søndergaard SA, Fløjgaard C, Ejrnæs R, Svenning J-C. Shifting Baselines and the Forgotten Giants: Integrating Megafauna into Plant Community Ecology. Oikos 2025: e11134.
  64. Soubry I, Guo X. Quantifying Woody Plant Encroachment in Grasslands: A Review on Remote Sensing Approaches. Canadian Journal of Remote Sensing 48: 337–378.
  65. Stein A, Gerstner K, Kreft H. Environmental Heterogeneity as a Universal Driver of Species Richness across Taxa, Biomes and Spatial Scales. Ecology Letters 17: 866–880.
    pubmed: 24751205
  66. Svenning JC. A Review of Natural Vegetation Openness in North‐Western Europe. Biological Conservation 104: 133–148.
  67. Svenning J-C, Buitenwerf R, E le Roux. Trophic Rewilding as a Restoration Approach under Emerging Novel Biosphere Conditions. Current Biology 34: R435–R451.
    pubmed: 38714176
  68. Svenning J-C, Lemoine RT, Bergman J, Buitenwerf R, Le Roux E, Lundgren E, Mungi N, Pedersen R Ø. The Late‐Quaternary Megafauna Extinctions: Patterns, Causes, Ecological Consequences and Implications for Ecosystem Management in the Anthropocene. Cambridge Prisms: Extinction 2: e5.
    pmc: PMC11895740pubmed: 40078803
  69. Svenning JC, Pedersen PBM, Donlan CJ, Ejrnæs R, Faurby S, Galetti M, Hansen DM. Science for a Wilder Anthropocene: Synthesis and Future Directions for Trophic Rewilding Research. Proceedings of the National Academy of Sciences of the United States of America 113: 898–906.
    pmc: PMC4743824pubmed: 26504218
  70. Tanentzap AJ, Daykin G, Fennell T, Hearne E, Wilkinson M, Carey PD, Woodcock BA, Heard MS. Trade‐Offs between Passive and Trophic Rewilding for Biodiversity and Ecosystem Functioning. Biological Conservation 281: 110005.
  71. Thomassen EE, Sigsgaard EE, Jensen MR, Olsen K, Hansen MDD, Svenning J-C, Thomsen PF. Contrasting Seasonal Patterns in Diet and Dung‐Associated Invertebrates of Feral Cattle and Horses in a Rewilding Area. Molecular Ecology 32: 2071–2091.
    pubmed: 36744391
  72. Timmermann A, Damgaard C, Strandberg MT, Svenning J-C. Pervasive Early 21st‐Century Vegetation Changes across Danish Semi‐Natural Ecosystems: More Losers than Winners and a Shift towards Competitive, Tall‐Growing Species. Journal of Applied Ecology 52: 21–30.
  73. Trepel J, le Roux E, Abraham AJ, Buitenwerf R, Kamp J, Kristensen JA, Tietje M, Lundgren EJ, Svenning J-C. Meta‐Analysis Shows that Wild Large Herbivores Shape Ecosystem Properties and Promote Spatial Heterogeneity. Nature Ecology & Evolution 8: 705–716.
    pubmed: 38337048
  74. Van Laer E, Moons CPH, Ampe B, Sonck B, Vangeyte J, Tuyttens F. Summertime Use of Natural Versus Artificial Shelter by Cattle in Nature Reserves. Animal Welfare 24: 345–356.
  75. Vera FWM, Bakker ES, Olff H. Large Herbivores: Missing Partners of Western European Light‐Demanding Tree and Shrub Species?. In Large Herbivore Ecology, Ecosystem Dynamics and Conservation, edited by Danell K.. 203–231. Cambridge: Cambridge University Press.
  76. Wang L, Cromsigt JPGM, Buitenwerf R, Lundgren EJ, Li W, Bakker ES, Svenning J-C. Tree Cover and its Heterogeneity in Natural Ecosystems Is Linked to Large Herbivore Biomass Globally. One Earth 6: 1759–1770.
  77. Zhang M, Yuan X, Zeng Z, Pan M, Wu P, Xiao J, Keenan TF. A Pronounced Decline in Northern Vegetation Resistance to Flash Droughts from 2001 to 2022. Nature Communications 16: 2984.
    pmc: PMC11947216pubmed: 40140384
  78. Zielke L, Wrage-Mönnig N, Müller J, Neumann C. Implications of Spatial Habitat Diversity on Diet Selection of European Bison and Przewalski's Horses in a Rewilding Area. Diversity 11: 63.

Citations

This article has been cited 0 times.
Subscribe to our newsletter
Join 100,129 horse owners like you who receive our email updates on horse health and nutrition.