Combination of herbivore removal and nitrogen deposition increases upland carbon storage.

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.

Calluna vulgaris-dominated upland heathland sequesters more CO2 annually than grass-dominated upland heathland.

It has been shown in many habitats worldwide, that a shift in vegetation composition between woody shrub and graminoid dominance can influence carbon (C) cycling. Due to land management practices and environmental change, UK upland heath vegetation has historically undergone shifts in dominance from the woody dwarf shrub Calluna vulgaris (Calluna) to species poor graminoid swards. The consequences of this for C sequestration are unknown. We compared annual net ecosystem exchange (NEE) of carbon dioxide (CO2) between building phase Calluna- and grass-dominated communities within three upland heaths in Scotland, measuring c. monthly over a year. Light and temperature response curves were generated, and the parameters derived were applied to continuous light and temperature data to extrapolate CO2 fluxes over the full year and generate estimates of annual CO2 sequestration for each vegetation type. Grass-dominated communities had higher ecosystem respiration rates than Calluna-dominated communities, attributed to graminoids having greater metabolic demands and producing more labile litter which decomposes readily. Both communities had similar gross primary productivity over the year; the net result being higher NEE within the Calluna-dominated than the grass-dominated community (-2.36 ± 0.23 and -1.78 ± 0.18 µmol CO2m(-2)s(-1) respectively). Modelled CO2 fluxes over a year showed both communities to be CO2 sinks. The Calluna-dominated community sequesters -3.45 ± 0.96 t C ha(-1)yr(-1), double that sequestered by the grass-dominated community at 1.61 ± 0.57 t C ha(-1)yr(-1). Potential rate of C sequestration by upland heath is comparable to that of woodland and the increase in total sequestration that could be gained from habitat restoration may equate to c. 60% of the annual UK C sink attributed to forest land management. National C sequestration by heathlands is also more than double that by peatlands. Management of graminoid-dominated upland heath should promote Calluna re-establishment, thus providing a C benefit in addition to benefits to biodiversity, grazing and sporting interests.