Temperate deciduous shrub phenology: the overlooked forest layer.

Temperate deciduous shrub phenology plays a pivotal role in forest ecology by regulating the timing of suitable habitat and food of a range of organisms as well as influencing the timing and duration of the carbon uptake period especially in early spring and late autumn when trees are leafless. However, given the overwhelming influence of canopy trees on forest ecosystem functioning, shrubs are often ignored in ecosystem modeling. Isolating the shrub contribution to C flux or satellite-derived forest phenology is challenging. In addition, since shrubs are more likely to be invasive than trees, future changes to shrub species composition are likely, with consequent implications for both over- and understory species composition and ecosystem functioning. Surprisingly, given their multifaceted role, our review revealed that studies on temperate deciduous shrub phenology are limited with the majority focusing on managing invasive shrubs in USA forests. In addition, results of some studies using a large number of species from a range of geographical locations suggested that, in general, invasive shrubs leafed out earlier and retained leaves longer than native species. However, this may not be directly applicable to local conditions with a smaller range of locally adapted species. Therefore, in order to fully understand the role of shrub phenology in temperate deciduous forests, in terms of invasive species, response to climate change and subsequent influence on C balance it will be necessary to establish phenological monitoring sites in which both tree and shrub phenology are recorded concurrently across a range of geographical locations.