Drought is a major environmental constrain affecting plant performance and survival, particularly in Mediterranean ecosystems. Terpenoids may play a protective role under these conditions, however, observations of drought effects on plant terpenoid emissions are controversial ranging from decreased emissions to unaffected or increased release of terpenoids. In the present study we investigated terpenoid emissions of cork oak (Quercus suber) and gum rockrose (Cistus ladanifer) in response to summer drought stress in 2017. Pre-dawn leaf water potential (ΨPD) decreased from -0.64 to -1.72 MPa in Q. suber and from -1.69 to -4.05 MPa in C. ladanifer, indicating a transition from mild to severe drought along summer. Total terpenoid emissions decreased with drought, but differed significantly between species (p < 0.001) and in response to ΨPD, air temperature and assimilation rates. C. ladanifer emitted a large variety of >75 compounds comprising monoterpenes, sesquiterpenes and even diterpenes, which strongly decreased from 1.37 ± 0.23 μg g-1h-1 to 0.40 ± 0.08 μg g-1h-1 (p < 0.001) in response to drought. Total emission rates were positively correlated to air temperature (p < 0.001). C. ladanifer behavior points toward terpenoid leaf storage depletion and reduced substrate availability for terpenoid synthesis with increasing drought, most likely accelerated by high air temperatures. Q. suber emitted mainly monoterpenes and emissions declined significantly from June (0.50 ± 0.08 μg g-1h-1) to August (0.29 ± 0.02 μg g-1h-1) (p < 0.01). Emission rates were weakly correlated with net assimilation rates (R2 = 0.19, p < 0.001), but did not respond strongly to ΨPD and air temperature. Early onset of drought in 2017 most likely reduced plant metabolism in Q. suber, resulting in diminished, but stable terpenoid fluxes. Calculation of standard emission factors (at 30°C) revealed contrasting emission patterns of decreasing, unaffected, or increasing fluxes of single terpenoid compounds. Unaffected or drought-enhanced emissions of compounds such as α-pinene, camphene or manoyl oxide may point toward a specific role of these terpenoids in abiotic stress adaptation. In conclusion, these results suggest a strong negative, but species- and compound-specific effect of severe drought on terpenoid fluxes in Mediterranean ecosystems.

Mediterranean vegetation emits a wide range of biogenic volatile organic compounds (BVOCs) among which isoprenoids present quantitatively the most important compound class. Here, we investigated the isoprenoid emission from two Mediterranean Cistaceae shrubs, Halimium halimifolium and Cistus ladanifer, under controlled and natural conditions, respectively. For the first time, diurnal emission patterns of the diterpene kaurene were detected in real-time by Proton-Transfer-Reaction-Time-of-Flight-Mass-Spectrometer. Kaurene emissions were strongly variable among H. halimifolium plants, ranging from 0.01 ± 0.003 to 0.06 ± 0.01 nmol m−2 s−1 in low and high emitting individuals, respectively. They were in the same order of magnitude as monoterpene (0.01 ± 0.01 to 0.11 ± 0.04 nmol m−2 s−1) and sesquiterpene (0.01 ± 0.01 to 0.52 nmol m−2 s−1) emission rates. Comparable range and variability was found for C. ladanifer under natural conditions. Labelling with 13C-pyruvate suggested that emitted kaurene was not derived from de novo biosynthesis. The high kaurene content in leaves, the weak relationship with ecophysiological parameters and the tendency of higher emissions with increasing temperatures in the field indicate an emission from storage pools. This study highlights significant emissions of kaurene from two Mediterranean shrub species, indicating that the release of diterpenes into the atmosphere should probably deserve more attention in the future.

 Interaction effects of different stressors, such as extreme drought and plant invasion, can have detrimental effects on ecosystem functioning and recovery after drought. With ongoing climate change and increasing plant invasion, there is an urgent need to predict the short-and long-term interaction impacts of these stressors on ecosystems.  We established a combined precipitation exclusion and shrub invasion (Cistus ladanifer) experiment in a Mediterranean cork oak (Quercus suber) ecosystem with four treatments: 1) Q. suber control, 2) Q. suber with rain exclusion, 3) Q. suber invaded by shrubs, 4) Q. suber with rain exclusion and shrub invasion. As key parameter, we continuously measured ecosystem water fluxes.  In an average precipitation year, the interaction effects of both stressors were neutral.However, the combination of imposed drought and shrub invasion led to amplifying interaction effects during an extreme drought by strongly reducing tree transpiration.Contrarily, the imposed drought reduced the competitiveness of the shrubs in the following recovery period, which buffered the negative effects of shrub invasion on Q. suber.  Our results demonstrate the highly dynamic and non-linear effects of interacting stressors on ecosystems and urges for further investigations on biotic interactions in a context of climate change pressures.