Mechanisms and implications of a type IV functional response for short-term intake rate of dry matter in large mammalian herbivores.

The functional response (i.e. the relationship between consumers' intake rate and resource density) is central in plant-herbivore interactions. Its shape and the biological processes leading to it have significant implications for both foraging theory and ecology of grazing systems. A type IV functional response (i.e. dome-shaped relationship) of short-term intake rate of dry matter (intake while grazing) has rarely been reported for large herbivores and the conditions that can lead to it are poorly understood. We report a type IV functional response observed in heifers grazing monocultures of Cynodon sp. and Avena strigosa. The mechanisms and consequences of this type of functional response for grazed system dynamics are discussed. Intake rate was higher at intermediate than at short or tall sward heights in both grass species. The type IV functional response resulted from changes in bite mass instead of a longer time needed to encounter and process bites. Thus, the decrease of intake rate of dry matter in tall swards is not explained by a shift from process 3 (potential bites are concentrated and apparent) to process 2 (potential bites are apparent but dispersed, Spalinger & Hobbs 1992). Bite mass was smaller in tall than in intermediate swards due to a reduction of bite volume possibly caused by the greater proportion of stem and sheath acting as a physical barrier to bite formation. It is generally accepted that potential bites are abundant and apparent in most grassland and meadow systems, as they were in the present experiments. Therefore, a type IV response of intake rate not directly related to digestive constraints may determine the dynamics of intake and defoliation under a much larger set of conditions than previously thought. These results have implications for foraging theory and stability of grazing systems. For example, if animals prefer patches of intermediate stature that yield the highest intake rate, grazing should lead to the widely observed bimodal distribution of plant mass per unit area, even when tall patches are not of significantly lower digestive quality than the pasture average.

Diet Switching by Mammalian Herbivores in Response to Exotic Grass Invasion.

Invasion by exotic grasses is a severe threat to the integrity of grassland ecosystems all over the world. Because grasslands are typically grazed by livestock and wildlife, the invasion is a community process modulated by herbivory. We hypothesized that the invasion of native South American grasslands by Eragrostis plana Nees, an exotic tussock-forming grass from Africa, could be deterred by grazing if grazers switched dietary preferences and included the invasive grass as a large proportion of their diets. Bos taurus (heifers) and Ovis aries (ewes) grazed plots with varying degrees of invasion by E. plana in a replicated manipulative experiment. Animal positions and species grazed were observed every minute in 45-min grazing session. Proportion of bites and steps in and out of E. plana tussocks were measured and used to calculate several indices of selectivity. Both heifers and ewes exhibited increasing probability of grazing E. plana as the proportion of area covered by tussocks increased, but they behaved differently. In agreement with expectations based on the allometry of dietary preferences and morphology, ewes consumed a low proportion of E. plana, except in areas that had more than 90% E. plana cover. Heifers consumed proportionally more E. plana than ewes. Contrary to our hypothesis, herbivores did not exhibit dietary switching towards the invasive grass. Moreover, they exhibited avoidance of the invasive grass and preference for short-statured native species, both of which should tend to enhance invasion. Unless invasive plants are highly palatable to livestock, the effect of grazing to deter the invasion is limited, due to the inherent avoidance of the invasive grass by the main grazers in the ecosystem, particularly sheep.