Not all temperate deciduous trees are leafless in winter: The curious case of marcescence

Abstract

AbstractTemperate deciduous forests by definition include a large proportion of woody species that shed their leaves each autumn and are completely leafless during winter months. Leaf senescence in deciduous trees is an active, complex process typically involving the physiological formation of an abscission layer causing the petiole to mechanically detach from the branch. However, several deciduous species retain all or some senesced leaves on branches through much of winter, a phenomenon called leaf marcescence. Marcescence has long fascinated botanists, including Pehr Kalm as early as 1749. Yet, surprisingly little research has been done to date. Here, we review and explore patterns and mechanisms of leaf marcescence in temperate forests, bringing together six nonmutually exclusive but separately proposed hypotheses: (1) Marcescence has no adaptive function but rather an evolutionary byproduct; (2) Marcescent leaves deter winter browsing herbivores; (3) Leaf retention through winter improves nutrient resorption during autumn senescence; (4) Prolonged leaf shedding into spring minimizes nutrient leaching and promotes decomposition; (5) Marcescent leaves protect overwintering buds from frost or desiccation; and (6) Marcescent canopies provide winter cover for animals (including insects, birds, bats), thereby affecting plant nutrient availability via excrement. No hypothesis has complete support and few tests of multiple hypotheses have been done. It is likely that any adaptive value of marcescence is species and context dependent. Despite increased interest in plant phenology and prevalence of this trait, much remains to be understood on the physiology, evolution, function, and ecological implications of leaf marcescence.