Negative effects of winter and spring warming on the regeneration of forest spring geophytes.

The climate is changing rapidly, provoking species to shift their ranges poleward and upslope. We currently lack a mechanistic understanding of the effect of warmer temperatures on plants, especially for seasonally distinct patterns. Spring geophytes are emblematic forest plants that have a short aboveground lifecycle in the first half of the year and are thus particularly sensitive to winter and spring warming. We set up a warming experiment with separate and combined winter and spring warming on seedlings of three European spring geophytes: Anemone nemorosa, Hyacinthoides non-scripta and Ornithogalum pyrenaicum. Seedling emergence and plant height were recorded at the end of winter and spring treatment, when also biomass of the root, shoot and storage organ was determined. We found negative effects of combined winter and spring warming on seedling emergence. The weight of the storage organ proved to be the best indicator of seedling performance and was negatively affected by separate winter warming in Anemone and by spring warming in Hyacinthoides. Successful seedling emergence was jeopardized by the absence of a cold period, while seedling performance seemed to be negatively influenced directly by higher temperatures through a phenological shift. Our findings confirm that warmer winter and spring temperatures could hamper regeneration of spring geophytes.

Forest understorey plant responses to long-term experimental warming, light and nitrogen addition.

Climate change, eutrophication and intensified forest management are affecting forest understorey plants, a major component of forest biodiversity. The main impacts of these drivers have often been studied, but we lack a good understanding of how key understorey species are affected by potential interactive effects of these drivers and which species drive community changes. Here we assessed the responses of 15 species occurring in the understorey of a deciduous temperate forest to experimental warming, light addition and enhanced nitrogen inputs in permanent plots surveyed for 9 years. We analysed vegetation cover and key functional traits (plant height, specific leaf area and reproductive traits) at the species level and identified the species driving community change with principal response curves (PRC). Light addition and warming, and to a lesser extent also nitrogen addition, had profound effects on cover and functional traits. Many species showed directional change over time, and this change can either be strengthened or weakened by treatments, indicating the importance of long-term monitoring. Against expectations, we observed few interactions between treatments. Species responses to treatments were related to ecological strategies (generalists versus forest specialist). Generalists, such as Rubus fruticosus, benefitted from the warming and light treatments and outcompeted forest specialists. This might ultimately lead to biotic homogenization. Since the treatment effects of light and warming were additive, keeping the canopy closed will only mitigate, but not stop, the effects of global warming on the forest understorey plants.

Individualistic responses of forest herb traits to environmental change.

Intraspecific trait variation (ITV; i.e. variability in mean and/or distribution of plant attribute values within species) can occur in response to multiple drivers. Environmental change and land-use legacies could directly alter trait values within species but could also affect them indirectly through changes in vegetation cover. Increasing variability in environmental conditions could lead to more ITV, but responses might differ among species. Disentangling these drivers on ITV is necessary to accurately predict plant community responses to global change. We planted herb communities into forest soils with and without a recent history of agriculture. Soils were collected across temperate European regions, while the 15 selected herb species had different colonizing abilities and affinities to forest habitat. These mesocosms (384) were exposed to two-level full-factorial treatments of warming, nitrogen addition and illumination. We measured plant height and specific leaf area (SLA). For the majority of species, mean plant height increased as vegetation cover increased in response to light addition, warming and agricultural legacy. The coefficient of variation (CV) for height was larger in fast-colonizing species. Mean SLA for vernal species increased with warming, while light addition generally decreased mean SLA for shade-tolerant species. Interactions between treatments were not important predictors. Environmental change treatments influenced ITV, either via increasing vegetation cover or by affecting trait values directly. Species' ITV was individualistic, i.e. species responded to different single resource and condition manipulations that benefited their growth in the short term. These individual responses could be important for altered community organization after a prolonged period.