Environmental data include long-term mean annual temperature (MAT) and precipitation (MAP) based on both older [63] and recent (unpublished weather station data) sources; this climatic regime has been broadly stable for the last 3000–5000 years [64]. Soil values are 0–5 cm means±s.d. of the ‘native’ cores sampled from upper and lower sites (N = 24). Bulk density, carbon and nitrogen, particulate organic matter (POM) C, and POM N all differed significantly (P<0.001) between the lower and upper sites. Soil cores were taken from areas of the two sites dominated by Poa spp.