Effects of browsing and grazing on cyclic succession in nutrient‐limited ecosystems

. This paper deals with browsing and grazing as forces driving cyclic succession. Between 1989 and 1994 reciprocal transitions between the dwarf shrub Calluna vulgaris and the grass Deschampsia flexuosa were monitored in permanent plots in a cattle grazed grass‐rich Dutch heathland on podsolic soils in which tree encroachment was prevented. Heather beetles killed Calluna in four of the nine plots during 1991/1992. The monitoring revealed reciprocal transitions and cycles between Calluna and Deschampsia on a subplot scale. Beetles and cattle had additional and complementary effects on the two competing species. Defoliation by beetles and trampling by cattle‐killed Calluna and favoured grass invasion. Grazing and gap creation by cattle in Deschampsia favoured the establishment and recovery of Calluna. Analysis of the causal mechanisms suggests that indirect, resource‐mediated herbivory effects may be as important for the replacement processes as direct effects of defoliation and trampling. Herbivory created differential light and nutrient levels in Calluna and Deschampsia gaps. Grazing and browsing improved the resource‐capturing abilities of Calluna and its resistance to herbivory and abiotic disturbances. The emerged Calluna‐Deschampsia cycle and its driving forces are summarized in a conceptual triangular resource‐mediated successional grazing cycle (RSGC) model, a limit cycle involving herbivore‐plant‐plant resource interactions. It offers a deterministic equilibrium model as alternative for stochastic transitions between the meta‐stable states with dominance of Calluna and Deschampsia respectively. The validity range of the RSGC model and its management implications are briefly discussed.