Forest understorey flowering phenology responses to experimental warming and illumination.

Species are altering their phenology to track warming temperatures. In forests, understorey plants experience tree canopy shading resulting in light and temperature conditions, which strongly deviate from open habitats. Yet, little is known about understorey phenology responses to forest microclimates. We recorded flowering onset, peak, end and duration of 10 temperate forest understorey plant species in two mesocosm experiments to understand how phenology is affected by sub-canopy warming and how this response is modulated by illumination, which is related to canopy change. Furthermore, we investigated whether phenological sensitivities can be explained by species' characteristics, such as thermal niche. We found a mean advance of flowering onset of 7.1 d per 1°C warming, more than previously reported in studies not accounting for microclimatic buffering. Warm-adapted species exhibited greater advances. Temperature sensitivity did not differ between early- and later-flowering species. Experimental illumination did not significantly affect species' phenological temperature sensitivities, but slightly delayed flowering phenology independent from warming. Our study suggests that integrating sub-canopy temperature and light availability will help us better understand future understorey phenology responses. Climate warming together with intensifying canopy disturbances will continue to drive phenological shifts and potentially disrupt understorey communities, thereby affecting forest biodiversity and functioning.

Forest floor environment overrules global change treatment effects on understorey communities in a mesocosm experiment.

Light availability profoundly influences plant communities, especially below dense tree canopies in forests. Canopy disturbances, altering forest floor light conditions, together with other environmental changes such as climate change, nitrogen deposition and legacy effects from previous land-use will simultaneously impact forest understorey communities. Yet, knowledge on the individual effects of these drivers and their potential interactions remains scarce. Here we performed a forest mesocosm experiment to assess the influence of warming, illumination (simulating canopy opening), nitrogen deposition and soil land-use history (comparing ancient and post-agricultural forest soil) on understorey community composition trajectories over a 7-year period. Strikingly, understorey communities primarily evolved in response to the deeply shaded ambient forest conditions, with experimental treatments exerting only secondary influences. The overruling trajectory steered all mesocosms towards slow-colonizing forest specialist communities dominated by spring geophytes with lower nutrient-demand. The illumination treatment and, to a lesser extent, warming and agricultural land-use legacy slowed down this trend by advancing fast-growing resource-acquisitive generalist species. Warm ambient temperatures induced thermophilization of plant communities in all treatments, including control plots, towards higher dominance of warm-adapted species. Nitrogen addition accelerated this thermophilization process and increased the community light-demand signature. Land-use legacy effects were limited in our study. Our findings underscore the essential role of limited light availability in preserving forest specialists in understorey communities and highlight the importance of maintaining a dense canopy cover to attenuate global change impacts. It is crucial to integrate this knowledge in forest management adaptation to global change, particularly in the face of increasing demands for wood and wood products and intensified natural canopy disturbances.

Combining multiple investigative approaches to unravel functional responses to global change in the understorey of temperate forests.

Plant communities are being exposed to changing environmental conditions all around the globe, leading to alterations in plant diversity, community composition, and ecosystem functioning. For herbaceous understorey communities in temperate forests, responses to global change are postulated to be complex, due to the presence of a tree layer that modulates understorey responses to external pressures such as climate change and changes in atmospheric nitrogen deposition rates. Multiple investigative approaches have been put forward as tools to detect, quantify and predict understorey responses to these global-change drivers, including, among others, distributed resurvey studies and manipulative experiments. These investigative approaches are generally designed and reported upon in isolation, while integration across investigative approaches is rarely considered. In this study, we integrate three investigative approaches (two complementary resurvey approaches and one experimental approach) to investigate how climate warming and changes in nitrogen deposition affect the functional composition of the understorey and how functional responses in the understorey are modulated by canopy disturbance, that is, changes in overstorey canopy openness over time. Our resurvey data reveal that most changes in understorey functional characteristics represent responses to changes in canopy openness with shifts in macroclimate temperature and aerial nitrogen deposition playing secondary roles. Contrary to expectations, we found little evidence that these drivers interact. In addition, experimental findings deviated from the observational findings, suggesting that the forces driving understorey change at the regional scale differ from those driving change at the forest floor (i.e., the experimental treatments). Our study demonstrates that different approaches need to be integrated to acquire a full picture of how understorey communities respond to global change.