Earthworms as drivers of the competition between grasses and legumes

Abstract

Grasses and legumes are grown together worldwide to improve total herbage yield and the quality of forage, however, the causes of population oscillations of grasses and legumes are poorly understood. Especially in grasslands, earthworms are among the most important detritivore animals functioning as ecosystem engineers, playing a key role in nutrient cycling and affecting plant nutrition and growth. The objectives of the present greenhouse experiment were to quantify the effects of earthworms on grass–legume competition in model grassland systems at two harvesting dates – simulating the widespread biannual mowing regime in Central European grasslands. The presence of earthworms increased the productivity of grasses and legumes after 6 weeks but only that of grasses after another 10 weeks. In mixed treatments, the presence of grasses and earthworms decreased legume shoot biomass, the amount of nitrogen (N) in shoot tissue and the number of legume flowerheads while the presence of legumes and earthworms increased the amount of N in grass shoots and the infestation of grasses with aphids. Analyses of 15N/ 14N ratios indicate that, compared to legumes, grasses more efficiently exploit soil mineral N and benefit from legume presence through reduced “intra-functional group” competition. In contrast to previous experiments, we found no evidence for N transfer from legumes to grasses. However, legume presence improved total herbage and N yield. Earthworms likely modulate the competition between grasses and legumes by increasing soil N uptake by plants and thereby increasing the competitive strength of grasses. Earthworms function as essential driving agents of grass–legume associations by (I) increasing grass yield, (II) increasing the amount of N in grass hay, (III) increasing the infestation rate of grasses with aphids, and (IV) potentially reducing the attractiveness of grass–legume associations to pollinators.