Herbivores sculpt leaf traits differently in grasslands depending on life form and land-use histories.

Vertebrate and invertebrate herbivores alter plant communities directly by selectively consuming plant species; and indirectly by inducing morphological and physiological changes to plant traits that provide competitive or survivorship advantages to some life forms over others. Progressively excluding aboveground herbivore communities (ungulates, medium and small sized mammals, invertebrates) over five growing seasons, we explored how leaf morphology (specific leaf area or SLA) and nutrition (nitrogen, carbon, phosphorous, potassium, sodium, and calcium) of different plant life forms (forbs, legumes, grasses, sedges) correlated with their dominance. We experimented in two subalpine grassland types with different land-use histories: (1) heavily grazed, nutrient-rich, short-grass vegetation and (2) lightly grazed, lower nutrient tall-grass vegetation. We found differences in leaf traits between treatments where either all herbivores were excluded or all herbivores were present, showing the importance of considering the impacts of both vertebrates and invertebrates on the leaf traits of plant species. Life forms responses to the progressive exclusion of herbivores were captured by six possible combinations: (1) increased leaf size and resource use efficiency (leaf area/nutrients) where lower nutrient levels are invested in leaf construction, but a reduction in the number of leaves, for example, forbs in both vegetation types, (2) increased leaf size and resource use efficiency, for example, legumes in short grass, (3) increased leaf size but a reduction in the number of leaves, for example, legumes in the tall grass, (4) increased number of leaves produced and increased resource use efficiency, for example, grasses in the short grass, (5) increased resource use efficiency of leaves only, for example, grasses and sedges in the tall grass, and (6) no response in terms of leaf construction or dominance, for example, sedges in the short grass. Although we found multiple possible responses by life forms to progressive exclusion of herbivores, we also found some important generalities. Changes in leaf traits of legumes and grasses correlated with their increasing dominance in the short-grass vegetation and plants were more efficient at constructing photosynthetic tissue when herbivores are present with few exceptions. These results demonstrate that vertebrate and invertebrate herbivores are essential to maintain plant species richness and resource-use efficiency.