Making the best of the worst of times: traits underlying combined shade and drought tolerance of Ruscus aculeatus and Ruscus microglossum (Asparagaceae)

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Abstract
The genus Ruscus (Asparagaceae) consists of evergreen, woody monocot shrubs with modified photosynthetic stems (phylloclades) that occur in dry, shaded woodland areas of the Mediterranean Basin and southern Europe. The combined drought and shade tolerance of Ruscus species challenges the ‘trade-off model’, which suggests that plants can be either drought or shade adapted, but not both. To clarify the potential mechanisms that enable Ruscus species to survive in shaded environments prone to pronounced soil drought, we studied form–function relations based on a detailed trait survey for Ruscus aculeatus L. and Ruscus microglossum Bertol., focusing on gas exchange, hydraulics, morphology, anatomy, and nutrient and isotope composition. We then compared these trait values with published data for other species. R. aculeatus and R. microglossum exhibited numerous traits conferring drought and shade tolerance via reduced demand for resources in general and an ability to survive on stored water. Specific traits include thick phylloclades with low rates of maximum photosynthetic CO2 assimilation, low stomatal conductance to water vapour (gs), low respiration rate, low light compensation point, low shoot hydraulic conductance, low cuticular conductance, and substantial water storage tissue. Ruscus carbon isotope composition values of –33 ‰ were typical of an understory plant, but given the low gs could be associated with internal CO2 recycling. Ruscus appears to be a model for extreme dual adaptation, both physiologically and morphologically, enabling its occupation of shaded sites within drought prone regions across a wide geographical range, including extremely low resource understory sites.

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