Abstract

In recent years, expansion of native and exotic evergreen shrubs into forest understories has been documented worldwide. Dense shrub thickets may interfere with tree establishment, suppress herbaceous cover, and contribute substantially to total standing crop of leaf biomass. Expansion may occur because evergreen shrubs exploit seasonal variations in irradiance and temperature that are characteristic of temperate understory environments. We quantified leaf-level light environment and photosynthetic activity of three sympatric broadleaf evergreens (Ilex opaca, Kalmia latifolia, and Myrica cerifera) in a deciduous forest understory in Charles City County,Virginia,USAin order to understand seasonal intra- and interspecific ranges of broadleaf evergreen physiology. Two species (K. latifolia and M. cerifera) represent a diverse taxonomic range within broadleaf evergreens, and often form expansive thickets. We measured parameters related to canopy structure (e.g., bifurcation ratio, leaf angle) and photosynthetic performance (e.g., electron transport rate or ETR, chlorophyll content), to identify potential mechanisms facilitating expansion. ETR varied both seasonally and among species. In summer, M. cerifera ETR was nearly double that ofI. opaca or K. latifolia. Additionally, leaf temperature enhanced photosynthetic capacity of expansive species. Evergreen species, though capable of fixing carbon throughout the year, often exhibit slow growth rates and low physiological activity. Yet, we observed that the range of evergreen physiological activity may be broader than previously recognized. Furthermore, our results indicate potential for changes in composition and expansion of the evergreen shrub layer by species that exhibit structural and physiological mechanisms advantageous for future rises in temperature.

Highlights

  • Expansion of evergreen shrubs, both native and exotic, (e.g., Ligustrum robustum, Ligustrum sinense, Myrica faya, Rosa multiflora) into forest understories worldwide has been documented [1,2,3,4,5]

  • Myrica cerifera, a nitrogen-fixing, evergreen shrub, which occurs in open environments and forest understories, has received attention for forming dense, monospecific thickets that encroach into nearby communities [10,11,12]

  • We investigated the response of evergreen understory woody species to seasonal differences in light environment and temperature by 1) quantifying seasonal variations in leaf pigments, as related to light availability, 2) identifying structural traits associated with light capture and that were related to photosynthetic capacity, 3) comparing photosynthetic capacity across seasons, and 4) determining if expansive, thicket-forming species (K. latifolia and M. cerifera) display physiology reflective of expansive potential in the understory

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Summary

Introduction

Both native and exotic, (e.g., Ligustrum robustum, Ligustrum sinense, Myrica faya, Rosa multiflora) into forest understories worldwide has been documented [1,2,3,4,5]. Within the Southern Appalachian Mountains of North America, 2.5 million ha are covered by thickets of Kalmia latifolia and Rhodedendron maximum [6,9]. Myrica cerifera, a nitrogen-fixing, evergreen shrub, which occurs in open environments and forest understories, has received attention for forming dense, monospecific thickets that encroach into nearby communities [10,11,12]. Species that form dense thickets may provide insight for understanding what suites of characteristics and physiological responses lead to occurrence and expansion within the understory

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