Combination of herbivore removal and nitrogen deposition increases upland carbon storage

Stuart W Smith, David Johnson, Samuel L O Quin, Kyle Munro, Robin J Pakeman, René van der Wal, Sarah J Woodin

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14 Citations (Scopus)
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Ecosystem carbon (C) accrual and storage can be enhanced by removing large herbivores as well as by the fertilizing effect of atmospheric nitrogen (N) deposition. These drivers are unlikely to operate independently, yet their combined effect on aboveground and belowground C storage remains largely unexplored. We sampled inside and outside 19 upland grazing exclosures, established for up to 80 years, across an N deposition gradient (5-24 kg N ha(-1) yr(-1) ) and found that herbivore removal increased aboveground plant C stocks, particularly in moss, shrubs and litter. Soil C storage increased with atmospheric N deposition, and this was moderated by the presence or absence of herbivores. In exclosures receiving above 11 kg N ha(-1) year(-1) , herbivore removal resulted in increased soil C stocks. This effect was typically greater for exclosures dominated by dwarf shrubs (Calluna vulgaris) than by grasses (Molinia caerulea). The same pattern was observed for ecosystem C storage. We used our data to predict C storage for a scenario of removing all large herbivores from UK heathlands. Predictions were made considering herbivore removal only (ignoring N deposition) and the combined effects of herbivore removal and current N deposition rates. Predictions including N deposition resulted in a smaller increase in UK heathland C storage than predictions using herbivore removal only. This finding was driven by the fact that the majority of UK heathlands receive low N deposition rates at which herbivore removal has little effect on C storage. Our findings demonstrate the crucial link between herbivory by large mammals and atmospheric N deposition, and this interaction needs to be considered in models of biogeochemical cycling.

Original languageEnglish
Pages (from-to)3036-3048
Number of pages13
JournalGlobal Change Biology
Issue number8
Early online date30 Apr 2015
Publication statusPublished - Aug 2015

Bibliographical note

© 2015 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.

We thank Ruth Mitchell, Alison Hester, Bob Mardon, Eoghain Maclean, David Welch, National Trust for Scotland, Scottish Natural Heritage and the Woodland Trust for helping find appropriate exclosures and granting access permission. We thank Nick Littlewood and Antonio Lopez Nogueira for their assistance in the field and processing samples in the lab and Ron Smith and Tony Dore for providing N deposition data. SWS was funded by a BBSRC studentship.


  • Calluna vulgaris
  • exclosures
  • grazing
  • heathlands
  • Molinia caerulea
  • nitrogen deposition
  • plant litter
  • soil carbon


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