Balancing positive and negative plant interactions: how mosses structure vascular plant communities.

Our understanding of positive and negative plant interactions is primarily based on vascular plants, as is the prediction that facilitative effects dominate in harsh environments. It remains unclear whether this understanding is also applicable to moss-vascular plant interactions, which are likely to be influential in low-temperature environments with extensive moss ground cover such as boreal forest and arctic tundra. In a field experiment in high-arctic tundra, we investigated positive and negative impacts of the moss layer on vascular plants. Ramets of the shrub Salix polaris, herb Bistorta vivipara, grass Alopecurus borealis and rush Luzula confusa were transplanted into plots manipulated to contain bare soil, shallow moss (3 cm) and deep moss (6 cm) and harvested after three growing seasons. The moss layer had both positive and negative impacts upon vascular plant growth, the relative extent of which varied among vascular plant species. Deep moss cover reduced soil temperature and nitrogen availability, and this was reflected in reduced graminoid productivity. Shrub and herb biomass were greatest in shallow moss, where soil moisture also appeared to be highest. The relative importance of the mechanisms by which moss may influence vascular plants, through effects on soil temperature, moisture and nitrogen availability, was investigated in a phytotron growth experiment. Soil temperature, and not nutrient availability, determined Alopecurus growth, whereas Salix only responded to increased temperature if soil nitrogen was also increased. We propose a conceptual model showing the relative importance of positive and negative influences of the moss mat on vascular plants along a gradient of moss depth and illustrate species-specific outcomes. Our findings suggest that, through their strong influence on the soil environment, mat-forming mosses structure the composition of vascular plant communities. Thus, for plant interaction theory to be widely applicable to extreme environments such as the Arctic, growth forms other than vascular plants should be considered.

Herbivore impacts to the moss layer determine tundra ecosystem response to grazing and warming.

Herbivory and climate are key environmental drivers, shaping ecosystems at high latitudes. Here, we focus on how these two drivers act in concert, influencing the high arctic tundra. We aim to investigate mechanisms through which herbivory by geese influences vegetation and soil processes in tundra ecosystems under ambient and warmed conditions. To achieve this, two grazing treatments, clipping plus faecal additions and moss removal, were implemented in conjunction with passive warming. Our key finding was that, in many cases, the tundra ecosystem response was determined by treatment impacts on the moss layer. Moss removal reduced the remaining moss layer depth by 30% and increased peak grass biomass by 27%. These impacts were probably due to observed higher soil temperatures and decomposition rates associated with moss removal. The positive impact of moss removal on grass biomass was even greater with warming, further supporting this conclusion. In contrast, moss removal reduced dwarf shrub biomass possibly resulting from increased exposure to desiccating winds. An intact moss layer buffered the soil to increased air temperature and as a result there was no response of vascular plant productivity to warming over the course of this study. In fact, moss removal impacts on soil temperature were nearly double those of warming, suggesting that the moss layer is a key component in controlling soil conditions. The moss layer also absorbed nutrients from faeces, promoting moss growth. We conclude that both herbivory and warming influence this high arctic ecosystem but that herbivory is the stronger driver of the two. Disturbance to the moss layer resulted in a shift towards a more grass-dominated system with less abundant mosses and shrubs, a trend that was further enhanced by warming. Thus herbivore impacts to the moss layer are key to understanding arctic ecosystem response to grazing and warming.