Journal of Coastal Conservation

, Volume 21, Issue 3, pp 369–379 | Cite as

Behaviour of horses and cattle at two stocking densities in a coastal salt marsh

  • S . NolteEmail author
  • C. van der Weyde
  • P. Esselink
  • C. Smit
  • S. E. van Wieren
  • J. P. Bakker


Livestock grazing has been practiced in salt marshes in the Wadden Sea area since 600 B.C. Currently livestock grazing is also applied for conservation management. However, effects of such grazing management on salt marshes are likely to vary depending on the species of livestock and stocking density due to differences in the behaviour of the animals. Yet, little is known about the behaviour of different livestock species and stocking densities grazing in salt marshes. We studied the grazing behaviour of horses and cattle by focal observation in an experiment with four different grazing treatments on a coastal salt marsh. In all treatments we recorded diet choice, movement and grazing activity, and spatial distribution. Livestock species shared an overlap in diet choice. Yet, horses more often foraged on the short grass Puccinellia maritima, while the cattle diet contained a higher amount of Aster tripolium. Horses travelled longer distances per day and spent more time grazing than cattle. Spatial distribution of cattle was significantly clustered, while horses showed a random distribution utilizing the whole area. Animal behaviour differs between livestock species and stocking densities with respect to diet choice, activity and spatial distribution.


Diet choice Focal observation Livestock species Conservation management Semi-natural grassland 



We thank ‘It Fryske Gea’ for logistic support and for kindly permitting us to setup the experiment. Gerrit van de Leest and Johannes Westerhof maintained the fences and inspected the well-being of the animals in the experiment. We acknowledge Anne-Marie van den Driessche for performing the chemical analysis and Nina Bhola for statistical advice. Adriënne Verburg and José de Jaeger are thanked for performing a preliminary study which enabled us to plan this research project. Dick Visser kindly prepared the figures. Esther Chang is acknowledged for improving the English of this manuscript. Sophie Prache and two reviewers gave valuable comments on a previous version of this manuscript. This research was funded by the Waddenfonds (project WF200451).

Supplementary material

11852_2017_515_MOESM1_ESM.pdf (540 kb)
Supplementary 1 (PDF 540 kb)
11852_2017_515_MOESM2_ESM.pdf (79 kb)
Supplementary 2 (PDF 78 kb)
11852_2017_515_MOESM3_ESM.pdf (21 kb)
Supplementary 3 (PDF 20 kb)


  1. Abrahamse PA, Tamminga S, Dijkstra J (2009) Effect of daily movement of dairy cattle to fresh grass in morning or afternoon on intake, grazing behaviour, rumen fermentation and milk production. J Agric Sci 147:721–730. doi: 10.1017/S0021859609990153 CrossRefGoogle Scholar
  2. Andresen H, Bakker JP, Brongers M et al (1990) Long-term changes of salt marsh communities by cattle grazing. Vegetatio 89:137–148CrossRefGoogle Scholar
  3. Augustine D, McNaughton S (1998) Ungulate effects on the functional species composition of plant communities: herbivore selectivity and plant tolerance. J Wildl Manag 62:1165–1183CrossRefGoogle Scholar
  4. Bakker JP, Leeuw J, Dijkema KS et al (1993) Salt marshes along the coast of The Netherlands. Hydrobiologia 265:73–95. doi: 10.1007/BF00007263 CrossRefGoogle Scholar
  5. Bakker JP, Bos D, De Vries Y (2003) To graze or not to graze: that is the question. In: Wolff W, Essink K, Van Leeuwe MA (eds) Challenges to the Wadden Sea area. (vol. 10, pp 67–87), Proceedings International Wadden Sea Symposium, GroningenGoogle Scholar
  6. Bates D, Mächler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67(1):1–48. doi: 10.18637/jss.v067.i01
  7. Berg G, Esselink P, Groeneweg M, Kiehl K (1997) Micropatterns in Festuca rubra-dominated salt-marsh vegetation induced by sheep grazing. Plant Ecol 132:1–14CrossRefGoogle Scholar
  8. Boyd LE, Carbonaro DA, Houpt KA (1988) The 24-hour time budget of Przewalski horses. Appl Anim Behav Sci 21:5–17. doi: 10.1016/0168-1591(88)90098-6 CrossRefGoogle Scholar
  9. ter Braak CJF, Šmilauer P (2002) CANOCO reference manual and CanoDraw for Windows user’s guide: software for canonical community ordination (version 4.5). Microcomputer power, IthacaGoogle Scholar
  10. Burnham KP, Anderson DR (2002) Model selection and Multimodel inference: a practical information-theoretic approach, 2nd edn. Springer-Verlag, New YorkGoogle Scholar
  11. Cornelissen P, Vulink JT (2015) Density-dependent diet selection and body condition of cattle and horses in heterogeneous landscapes. Appl Anim Behav Sci 163:28–38. doi: 10.1016/j.applanim.2014.12.008 CrossRefGoogle Scholar
  12. Crawley MJ (1983) Herbivory - the dynamics of animal-plant interactions (studies in Ecology 10). Blackwell Scientific, OxfordGoogle Scholar
  13. Dale AJ, Mayne CS, Laidlaw AS, Ferris CP (2008) Effect of altering the grazing interval on growth and utilization of grass herbage and performance of dairy cows under rotational grazing. Grass Forage Sci 63:257–269CrossRefGoogle Scholar
  14. Development Core Team R (2013) R: a language and environment for statistical computing. R Foundation for Statistical Computing, ViennaGoogle Scholar
  15. Dixon PM (2006) Nearest neighbor methods. Encycl environmetrics. doi: 10.1002/9780470057339.van007
  16. Dumont B, Garel JP, Ginane C et al (2007a) Effect of cattle grazing a species-rich mountain pasture under different stocking rates on the dynamics of diet selection and sward structure. Animal 1:1042–1052. doi: 10.1017/S1751731107000250 CrossRefGoogle Scholar
  17. Dumont B, Rook AJ, Coran C, Röver KU (2007b) Effects of livestock breed and grazing intensity on biodiversity and production in grazing systems. 2. Diet selection. Grass Forage Sci 62:159–171CrossRefGoogle Scholar
  18. Dumont B, Rossignol N, Loucougaray G et al (2012) When does grazing generate stable vegetation patterns in temperate pastures? Agric Ecosyst Environ 153:50–56. doi: 10.1016/j.agee. 2012.03.003 CrossRefGoogle Scholar
  19. Duncan P, Foose TJ, Gordon IJ et al (1990) Comparative nutrient extraction from forages by grazing bovids and equids: a test of the nutritional model of equid / bovid competition and coexistence. Oecologia 84:411–418CrossRefGoogle Scholar
  20. Elizalde-Arellano C, Lopez-Vidal JC, Hernandez L et al (2012) Home range size and activity patterns of bobcats (Lynx rufus) in the southern part of their range in the Chihuahuan Desert, Mexico. Am Midl Nat 168:247–264CrossRefGoogle Scholar
  21. ESRI (2011) ArcGIS Desktop: Release 10. Environmental Systems Research Institute, Redlands, CAGoogle Scholar
  22. Esselink P, Zijlstra W, Dijkema KS, van Diggelen R (2000) The effects of decreased management on plant-species distribution patterns in a salt marsh nature reserve in the Wadden Sea. Biol Conserv 93:61–76CrossRefGoogle Scholar
  23. Esselink P, Fresco LFM, Dijkema KS (2002) Vegetation change in a man-made salt marsh affected by a reduction in both grazing and drainage. Appl Veg Sci 5:17–32. doi: 10.1658/1402-2001(2002)005[0017:VCIAMM]2.0.CO;2 Google Scholar
  24. Ferreira LMM, Celaya R, Benavides R et al (2013) Foraging behaviour of domestic herbivore species grazing on heathlands associated with improved pasture areas. Livest Sci 155:373–383. doi: 10.1016/j.livsci.2013.05.007 CrossRefGoogle Scholar
  25. Gibb MJ, Huckle CA, Nuthall R, Rook AJ (1999) The effect of physiological state (lactating or dry) and sward surface height on grazing behaviour and intake by dairy cows. Appl Anim Behav Sci 63:269–287. doi: 10.1016/S0168-1591(99)00014-3 CrossRefGoogle Scholar
  26. Gordon IJ (1989) Vegetation Community selection by ungulates on the isle of Rhum. II Vegetation Community Selection J Appl Ecol 26:53–64Google Scholar
  27. Hampson BA, Morton JM, Mills PC et al (2010) Monitoring distances travelled by horses using GPS tracking collars. Aust Vet J 88:176–181. doi: 10.1111/j.1751-0813.2010.00564.x CrossRefGoogle Scholar
  28. Jensen A (1985) The effect of cattle and sheep grazing on salt-marsh vegetation at Skallingen, Denmark. Vegetatio 60:37–48. doi: 10.1007/BF00053910 CrossRefGoogle Scholar
  29. Karmiris I, Platis PPD, Kazantzidis S, Papachristou TG (2011) Diet selection by domestic and wild herbivore species in a coastal Mediterranean wetland. Ann Zool Fennici 48:233–242CrossRefGoogle Scholar
  30. Kiehl K, Eischeid I, Gettner S, Walter J (1996) Impact of different sheep grazing intensities on salt marsh vegetation in northern Germany. J Veg Sci 7:99–106CrossRefGoogle Scholar
  31. Kiehl K, Esselink P, Gettner S, Bakker JP (2001) The impact of sheep grazing on net nitrogen mineralization rate in two temperate salt marshes. Plant Biol 3:553–560CrossRefGoogle Scholar
  32. Kiehl K, Schröder H, Stock M (2007) Long-term vegetation dynamics after land-use change in Wadden Sea salt marshes. Coastline Reports 7:17–24Google Scholar
  33. Kleyer M, Feddersen H, Bockholt R (2003) Secondary succession on a high salt marsh at different grazing intensities. J Coast Conserv 9:123–134CrossRefGoogle Scholar
  34. van Klink R, Nolte S, Mandema FS et al (2016) Effects of grazing management on biodiversity across trophic levels–the importance of livestock species and stocking density in salt marshes. Agric Ecosyst Environ 235:329–339. doi: 10.1016/j.agee.2016.11.001 CrossRefGoogle Scholar
  35. Krysl LJ, Hubbert ME, Sowell BF, et al (1984) Horses and Cattle Grazing in the Wyoming Red Desert .1. Food-Habits and Dietary Overlap. J Range Manag 37:72–76.Google Scholar
  36. Lellau TF, Liebezeit G (2001) Alkaloids, saponins and phenolic compounds in salt marsh plants from the lower Saxonian Wadden Sea. Senckenberg Marit 31:1–9. doi: 10.1007/BF03042831 CrossRefGoogle Scholar
  37. Mandema FS, Tinbergen JM, Ens BJ, Bakker JP (2013) Livestock grazing and trampling of birds’ nests: an experiment using artificial nests. J Coast Conserv 17:409–416. doi: 10.1007/s11852-013-0239-2 CrossRefGoogle Scholar
  38. Mandema FS, Tinbergen JM, Ens BJ, Bakker JP (2014) Spatial diversity in canopy height at redshank and oystercatcher Nest-sites in relation to livestock grazing. Ardea 101:105–112. doi: 10.5253/078.101.0205 CrossRefGoogle Scholar
  39. Menard C, Duncan P, Fleurance G et al (2002) Comparative foraging and nutrition of horses and cattle in European wetlands. J Appl Ecol 39:120–133CrossRefGoogle Scholar
  40. Metera E, Sakowski T, Sloniewski K, Romanowicz B (2010) Grazing as a tool to maintain biodiversity of grassland - a review. Anim Sci Pap Reports 28:315–334Google Scholar
  41. Meyer H, Fock H, Haase A et al (1995) Structure of the invertebrate Fauna in salt marshes of the Wadden Sea coast of Schleswig-Holstein influenced by sheep-grazing. Helgolander Meeresuntersuchungen 49:563–589. doi: 10.1007/BF02368383 CrossRefGoogle Scholar
  42. Nolte S, Esselink P, Bakker JP (2013) Flower production of Aster tripolium is affected by behavioral differences in livestock species and stocking densities: the role of activity and selectivity. Ecol Res 28:821–831CrossRefGoogle Scholar
  43. Nolte S, Esselink P, Smit C, Bakker JP (2014) Herbivore species and density affect vegetation-structure patchiness in salt marshes. Agric Ecosyst Environ 185:41–47. doi: 10.1016/j.agee.2013.12.010 CrossRefGoogle Scholar
  44. Nolte S, Esselink P, Bakker JP, Smit C (2015) Effects of livestock species and stocking density on accretion rates in grazed salt marshes. Estuar Coast Shelf Sci 152:109–115. doi: 10.1016/j.ecss.2014.11.012 CrossRefGoogle Scholar
  45. Norris K, Cook T, O’Dowd B, Durdin C (1997) The density of redshank Tringa totanus breeding on the salt-marshes of the wash in relation to habitat and its grazing management. J Appl Ecol 34:999–1013CrossRefGoogle Scholar
  46. Pepin D, Morellet N, Goulard M (2009) Seasonal and daily walking activity patterns of free-ranging adult red deer (Cervus elaphus) at the individual level. Eur J Wildl Res 55:479–486CrossRefGoogle Scholar
  47. Pinheiro J, Bates D, DebRoy S, Sarkar D, R Core Team (2017). nlme: linear and nonlinear mixed effects models. R package version 3.1–131.
  48. Provenza FD, Balph DF (1987) Diet learning by domestic ruminants: theory, evidence and practical implications. Appl Anim Behav Sci 18:211–232CrossRefGoogle Scholar
  49. Putman RJ, Fowler AD, Tout S (1991) Patterns of use of ancient grassland by cattle and horses and effects on vegetational composition and structure. Biol Conserv 56:329–347. doi: 10.1016/0006-3207(91)90065-H CrossRefGoogle Scholar
  50. Rickert C (2011) Microlepidoptera in salt marshes - life history, effects of grazing, and their suitability as ecological indicators. Suppl zu Faun ökologische Mitteilungen 37:5–124Google Scholar
  51. Rickert C, Fichtner A, van Klink R, Bakker JP (2012) α- and β-diversity in moth communities in salt marshes is driven by grazing management. Biol Conserv 146:24–31. doi: 10.1016/j.biocon.2011.11.024 CrossRefGoogle Scholar
  52. Rook AJ, Dumont B, Isselstein J et al (2004) Matching type of livestock to desired biodiversity outcomes in pastures – a review. Biol Conserv 119:137–150. doi: 10.1016/j.biocon.2003.11.010 CrossRefGoogle Scholar
  53. Rutter SM, Orr RJ, Yarrow NH, Champion RA (2004) Dietary preference of dairy cows grazing ryegrass and white clover. J Dairy Sci 87:1317–1324. doi: 10.3168/jds.S0022-0302(04)73281-6 CrossRefGoogle Scholar
  54. Schröder HK, Kiehl K, Stock M (2002) Directional and non-directional vegetation changes in a temperate salt marsh in relation to biotic and abiotic factors. Appl Veg Sci 5:33–44Google Scholar
  55. Silliman BR, Mozdzer T, Angelini C et al (2014) Livestock as potential biological control agent for an invasive wetland plant. PeerJ. doi: 10.7717/peerj.567 Google Scholar
  56. Stewart GB, Pullin AS (2008) The relative importance of grazing stock type and grazing intensity for conservation of mesotrophic “old meadow” pasture. J Nat Conserv 16:175–185. doi: 10.1016/j.jnc.2008.09.005 CrossRefGoogle Scholar
  57. Van Soest PJ, Robertson JB, Lewis BA (1991) Methods for Dietary fiber, neutral detergent fiber, and Nonstarch polysaccharides in relation to animal nutrition. J Dairy Sci 74:3583–3597CrossRefGoogle Scholar
  58. Veeneklaas RM, Dijkema KS, Hecker N, Bakker JP (2013) Spatio-temporal dynamics of the invasive plant species Elytrigia atherica on natural salt marshes. Appl Veg Sci 16:205–216CrossRefGoogle Scholar
  59. Vulink JT, Drost HJ (1991) A causal analysis of diet composition in free ranging cattle in reed-dominated vegetation. Oecologia 88:167–172CrossRefGoogle Scholar
  60. Westhoff V, Hobohm C, Schaminée JHJ (1993) Rote Liste der Pflanzengesellschaften des Naturraumes Wattenmeer unter Berücksichtigung der ungefährdeten Vegetationseinheiten. Tuexenia 13:109–140Google Scholar
  61. Zuur A, Ieno EN, Walker N, Saveliev AA, Smith GM (2009) Mixed effects models and extensions in ecology with R. Statistics for Biology and Health. Springer, New YorkGoogle Scholar

Copyright information

© Springer Science+Business Media Dordrecht 2017

Authors and Affiliations

  • S . Nolte
    • 1
    • 2
    Email author
  • C. van der Weyde
    • 3
  • P. Esselink
    • 1
    • 4
  • C. Smit
    • 1
  • S. E. van Wieren
    • 5
  • J. P. Bakker
    • 1
  1. 1.Conservation Ecology, Groningen Institute of Evolutionary Life Sciences GELIFESUniversity of GroningenGroningenThe Netherlands
  2. 2.Applied Plant Ecology, Biocenter Klein FlottbekUniversity of HamburgHamburgGermany
  3. 3.Van Hall Larenstein University of Applied SciencesLeeuwardenThe Netherlands
  4. 4.Puccimar, Ecological Research and ConsultancyVriesThe Netherlands
  5. 5.Resource Ecology GroupWageningen UniversityWageningenThe Netherlands

Personalised recommendations