Unpacking multi-trophic herbivore-grass-endophyte interactions: feedbacks across different scales in vegetation responses to Soay sheep herbivory.

Grazing can induce changes in both plant productivity and nutritional quality, which may subsequently influence herbivore carrying capacity. While research on Soay sheep (Ovis aries L.) dynamics on Hirta Island in the St. Kilda archipelago has elucidated the complexity of population drivers, including parasites, the role of herbivore-generated feedbacks as an intrinsic regulating factor remains unclear. The sheep lack large predators and every 3-9 years undergo population crashes (overcompensatory mortality). We investigated the effects of grazing on (1) sward productivity and (2) quality (toxicity) of the primary forage species, red fescue (Festuca rubra L.), which is highly infected by an alkaloid-synthesizing fungal endophyte. Grazing had a negative impact on both forage quantity and quality. At higher sheep densities, impacts on sward growth were magnified, resulting in a nonlinear relationship with plant productivity. Simultaneously, endophyte hyphal load (and by inference, toxicity) peaked close to the time of a crash. A greenhouse experiment showed that alkaloid concentration in F. rubra increased in response to artificial defoliation. We conclude that at high sheep densities, grazing-mediated reductions in productivity, together with sustained alkaloid production, are likely to influence sheep dynamics. Future research should consider the interactive effects of forage toxicity, quantity, and nutritional content.

Effect of Herbivory and Plant Species Replacement on Primary Production.

Grazing optimization occurs when herbivory increases primary production at low grazing intensities. In the case of simple plant-herbivore interactions, such an effect can result from recycling of a limiting nutrient. However, in more complex cases, herbivory can also lead to species replacement in plant communities, which in turn alters how primary production is affected by herbivory. Here we explore this issue using a model of a limiting nutrient cycle in an ecosystem with two plant species. We show that two major plant traits determine primary production at equilibrium: plant recycling efficiency (i.e., the fraction of the plant nutrient stock that stays within the ecosystem until it is returned to the nutrient pool in mineral form) and plant ability to deplete the soil mineral nutrient pool through consumption of this resource. In cases where sufficient time has occurred, grazing optimization requires that herbivory improve nutrient conservation in the system sufficiently. This condition sets a minimum threshold for herbivore nutrient recycling efficiency, the fraction of nutrient consumed by herbivores that is recycled within the ecosystem to the mineral nutrient pool. This threshold changes with plant community composition and herbivore preference and is, therefore, strongly affected by plant species replacement. The quantitative effects of these processes on grazing optimization are determined by both the recycling efficiencies and depletion abilities of the plant species. However, grazing optimization remains qualitatively possible even with plant species replacement.