Indirect short- and long-term effects of aboveground invertebrate and vertebrate herbivores on soil microarthropod communities.

Recognition is growing that besides ungulates, small vertebrate and invertebrate herbivores are important drivers of grassland functioning. Even though soil microarthropods play key roles in several soil processes, effects of herbivores-especially those of smaller body size-on their communities are not well understood. Therefore, we progressively excluded large, medium and small vertebrate and invertebrate herbivores for three growing seasons using size-selective fences in two vegetation types in subalpine grasslands; short-grass and tall-grass vegetation generated by high and low historical levels of ungulate grazing. Herbivore exclusions generally had few effects on microarthropod communities, but exclusion of all herbivore groups resulted in decreased total springtail and Poduromorpha richness compared with exclusion of only ungulates and medium-sized mammals, regardless of vegetation type. The tall-grass vegetation had a higher total springtail richness and mesostigmatid mite abundance than the short-grass vegetation and a different oribatid mite community composition. Although several biotic and abiotic variables differed between the exclusion treatments and vegetation types, effects on soil microarthropods were best explained by differences in nutrient and fibre content of the previous year's vegetation, a proxy for litter quality, and to a lesser extent soil temperature. After three growing seasons, smaller herbivores had a stronger impact on these functionally important soil microarthropod communities than large herbivores. Over longer time-scales, however, large grazers created two different vegetation types and thereby influenced microarthropod communities bottom-up, e.g. by altering resource quality. Hence, both short- and long-term consequences of herbivory affected the structure of the soil microarthropod community.

Aboveground vertebrate and invertebrate herbivore impact on net N mineralization in subalpine grasslands.

Aboveground herbivores have strong effects on grassland nitrogen (N) cycling. They can accelerate or slow down soil net N mineralization depending on ecosystem productivity and grazing intensity. Yet, most studies only consider either ungulates or invertebrate herbivores, but not the combined effect of several functionally different vertebrate and invertebrate herbivore species or guilds. We assessed how a diverse herbivore community affects net N mineralization in subalpine grasslands. By using size-selective fences, we progressively excluded large, medium, and small mammals, as well as invertebrates from two vegetation types, and assessed how the exclosure types (ET) affected net N mineralization. The two vegetation types differed in long-term management (centuries), forage quality, and grazing history and intensity. To gain a more mechanistic understanding of how herbivores affect net N mineralization, we linked mineralization to soil abiotic (temperature; moisture; NO3-, NH4+, and total inorganic N concentrations/pools; C, N, P concentrations; pH; bulk density), soil biotic (microbial biomass; abundance of collembolans, mites, and nematodes) and plant (shoot and root biomass; consumption; plant C, N, and fiber content; plant N pool) properties. Net N mineralization differed between ET, but not between vegetation types. Thus, short-term changes in herbivore community composition and, therefore, in grazing intensity had a stronger effect on net N mineralization than long-term management and grazing history. We found highest N mineralization values when only invertebrates were present, suggesting that mammals had a negative effect on net N mineralization. Of the variables included in our analyses, only mite abundance and aboveground plant biomass explained variation in net N mineralization among ET. Abundances of both mites and leaf-sucking invertebrates were positively correlated with aboveground plant biomass, and biomass increased with progressive exclusion. The negative impact of mammals on net N mineralization may be related partially to (1) differences in the amount of plant material (litter) returned to the belowground subsystem, which induced a positive bottom-up effect on mite abundance, and (2) alterations in the amount and/or distribution of dung, urine, and food waste. Thus, our results clearly show that short-term alterations of the aboveground herbivore community can strongly impact nutrient cycling within ecosystems independent of long-term management and grazing history.