Both bottom-up (e.g., nutrients) and top-down (e.g., herbivory) forces structure plant communities, but it remains unclear how they affect the relative importance of stochastic and deterministic processes in plant community assembly. Moreover, different-sized herbivores have been shown to have contrasting effects on community structure and function, but their effects on the processes governing community assembly (i.e., how they generate the impacts on structure) remain largely unknown. We evaluated the influence of bottom-up and top-down forces on the relative importance of deterministic and stochastic processes during plant community assembly. We used the data of a 7-yr factorial experiment manipulating nutrient availability (ambient and increased) and the presence of vertebrate herbivores (>1 kg) of different body size in a floodplain grassland in The Netherlands. We used a null model that describes a community composition expected by chance (i.e., stochastic assembly) and compared the plant community composition in the different treatments with this null model (the larger the difference, the more deterministically assembled). Our results showed that herbivore exclusion promoted a more stochastic plant community assembly, whereas increased nutrients played a relatively minor role in determining the relative importance of stochasticity in community assembly. Large herbivores facilitated intermediate-sized mammal herbivores, resulting in synergistic effects of enhanced grazing pressure and a more deterministic and convergent plant community assembly. We conclude that herbivores can act as strong deterministic forces during community assembly in natural systems. Our results also reveal that although large- and intermediate-sized mammal herbivores often have contrasting effects on many community and ecosystem properties, they can also synergistically homogenize plant communities.

Conservation Ecology Group Groningen Institute of Evolutionary Life Sciences University of Groningen P O Box 11103 9700 CC Groningen The Netherlands

Department of Aquatic Ecology Netherlands Institute of Ecology Droevendaalsesteeg 10 6708 PB Wageningen The Netherlands

Institute of Botany Czech Academy of Sciences Dukelská 135 37982 Tĭeboň Czech Republic

Laboratorio de Ecología Instituto de Investigaciones Marinas y Costeras CC 1260 Correo Central B7600WAG Mar del Plata Argentina

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