Precipitation-dependent flowering of Globularia alypum and Erica multiflora in Mediterranean shrubland under experimental drought and warming, and its inter-annual variability

Background and Aims

Relationships between autumn flowering, precipitation and temperature of plant species of Mediterranean coastal shrublands have been described, but not analysed experimentally. These relationships were analysed for two species of co-occurring, dominant, autumn-flowering shrubs, Globularia alypum and Erica multiflora, over 4 years and in experimentally generated drought and warming conditions. The aim was to improve predictions about the responses and adaptations of flowering of Mediterranean vegetation to climate change.

Methods

Beginning of anthesis and date of maximum flowering intensity (‘peak date’) were monitored over 4 years (2001–2004) on a garrigue land type in the noth-east of the Iberian Peninsula. Two experimental treatments were applied, increased temperature (+0·73°C) and reduced soil moisture (–17%) relative to untreated plots.

Key Results

Flowering of Globularia alypum and Erica multiflora differed greatly between years depending on the precipitation of the previous months and the date of the last substantial rainfall (>10 mm). Globularia alypum flowered once or twice (unimodal or bimodal) as the result of differences in the distribution and magnitude of precipitation in late-spring and summer (when floral buds develop). The drought treatment delayed and decreased flowering of Globularia alypum in 2001 and delayed flowering in 2002. Warming extended the period between the beginning of flowering and the end of the second peak for autumn flowering in 2001 and also increased peak intensity in 2002. Flowering of Erica multiflora was unaffected by either treatment.

Conclusions

Autumn flowering of Globularia alypum and Erica multiflora is more dependent on water availability than on temperature. Considerable inter-annual plasticity in the beginning of anthesis and peak date and on unimodal or bimodal flowering constitutes a ‘safe strategy’ for both species in relation to varying precipitation and temperature. However, severe changes in precipitation in spring and summer may severely affect flowering of Globularia alypum but not Erica multiflora, thus affecting development/structure of the ecosystem if such conditions persist.Key words: Globularia alypum, Erica multiflora, autumn flowering, drought, global warming, Mediterranean     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Few multiyear precipitation–reduction experiments find a shift in the productivity–precipitation relationship

Well‐defined productivity–precipitation relationships of ecosystems are needed as benchmarks for the validation of land models used for future projections. The productivity–precipitation relationship may be studied in two ways: the spatial approach relates differences in productivity to those in precipitation among sites along a precipitation gradient (the spatial fit, with a steeper slope); the temporal approach relates interannual productivity changes to variation in precipitation within sites (the temporal fits, with flatter slopes). Precipitation–reduction experiments in natural ecosystems represent a complement to the fits, because they can reduce precipitation below the natural range and are thus well suited to study potential effects of climate drying. Here, we analyse the effects of dry treatments in eleven multiyear precipitation–manipulation experiments, focusing on changes in the temporal fit. We expected that structural changes in the dry treatments would occur in some experiments, thereby reducing the intercept of the temporal fit and displacing the productivity–precipitation relationship downward the spatial fit. The majority of experiments (72%) showed that dry treatments did not alter the temporal fit. This implies that current temporal fits are to be preferred over the spatial fit to benchmark land‐model projections of productivity under future climate within the precipitation ranges covered by the experiments. Moreover, in two experiments, the intercept of the temporal fit unexpectedly increased due to mechanisms that reduced either water loss or nutrient loss. The expected decrease of the intercept was observed in only one experiment, and only when distinguishing between the late and the early phases of the experiment. This implies that we currently do not know at which precipitation–reduction level or at which experimental duration structural changes will start to alter ecosystem productivity. Our study highlights the need for experiments with multiple, including more extreme, dry treatments, to identify the precipitation boundaries within which the current temporal fits remain valid.     

Effects of long-term experimental night-time warming and drought on photosynthesis, <Emphasis Type="Italic">F</Emphasis>v/<Emphasis Type="Italic">F</Emphasis>m and stomatal conductance in the dominant species of a Mediterranean shrubland

We conducted a night-time warming and drought field experiment for 7 years (1999–2005) in a Mediterranean shrubland. We focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree Pinus halepensis L. and the final years to study the effects of the experimental night-time warming and drought on Fv/Fm, photosynthesis, and stomatal conductance. Warming treatment increased mean air temperature and mean soil temperature through the years by an average of 0.7 and 0.9°C respectively, and drought treatment reduced soil moisture through the years by an average of 19%. Warming tended to increase photosynthetic rates in E. multiflora, G. alypum and P. halepensis mostly in the cold seasons, when plants were more limited by temperature, as shown by the lowest values of Fv/Fm being detected in winter in the three studied species. A negative effect of warming was only detected for E. multiflora in summer 2003. Drought treatment generated different responses of net photosynthetic rates depending on the species, season and year. Stomatal conductance showed the same pattern as photosynthesis for the three studied species, displaying seasonal and inter-annual variability, although with an overall negative effect of drought for P. halepensis. Photosynthetic rates decreased significantly in the dry winter 2005 and spring 2005 in comparison to the same seasons of 2003 and 2004. There were positive correlations between the photosynthetic rates in different seasons for E. multiflora, G. alypum and P. halepensis and the soil moisture of the week prior to measurements. The great variation in the photosynthetic rates was thus explained in a significant part by soil moisture levels. The lowest Fv/Fm values usually corresponded with lowest stomatal conductances suggesting that drought stress could be associated to stress by low temperatures in winter.     

Temperature Dependence of Soil Respiration Modulated by Thresholds in Soil Water Availability Across European Shrubland Ecosystems

Soil respiration (SR) is a major component of the global carbon cycle and plays a fundamental role in ecosystem feedback to climate change. Empirical modelling is an essential tool for predicting ecosystem responses to environmental change, and also provides important data for calibrating and corroborating process-based models. In this study, we evaluated the performance of three empirical temperature–SR response functions (exponential, Lloyd–Taylor and Gaussian) at seven shrublands located within three climatic regions (Atlantic, Mediterranean and Continental) across Europe. We investigated the performance of SR models by including the interaction between soil moisture and soil temperature. We found that the best fit for the temperature functions depended on the site-specific climatic conditions. Including soil moisture, we identified thresholds in the three different response functions that improved the model fit in all cases. The direct soil moisture effect on SR, however, was weak at the annual time scale. We conclude that the exponential soil temperature function may only be a good predictor for SR in a narrow temperature range, and that extrapolating predictions for future climate based on this function should be treated with caution as modelled outputs may underestimate SR. The addition of soil moisture thresholds improved the model fit at all sites, but had a far greater ecological significance in the wet Atlantic shrubland where a fundamental change in the soil CO₂ efflux would likely have an impact on the whole carbon budget.     

Net ecosystem exchange and whole plant isoprenoid emissions by a mediterranean shrubland exposed to experimental climate change

We tested the effect of forecasted soil drought and warming climate conditions for the next decades on emission rates of isoprenoids by mediterranean shrublands. We measured isoprenoid emissions by whole dominant mediterranean woody plants (Erica multiflora L. and Globularia alypum L.) inhabiting the studied shrublands. Monoterpene emissions were detected in both species, but isoprene was emitted only by E. multiflora. Maximum emission rates were found during the hottest periods (except for G. alypum, in which they occurred in autumn), and minimum emission rates in winter in E. multiflora. Terpene emission rates ranged from 0.08 μg/(g dry wt h) in winter in E. multiflora to 8.8 μg/(g dry wt h) in G. alypum in autumn. In E. multiflora, the terpene emission rates decreased in response to soil drought only in summer, but increased in response to warming in spring and autumn. Isoprene emissions ranged from 0.1 μg/(g dry wt h) in spring to 4.4 μg/(g dry wt h) in summer. The effect of the treatments was only detected in autumn when soil drought and warming had a negative effect on isoprene emission rates. These data might improve our knowledge of isoprenoid emissions at the canopy level and in response to climate change, soil drought, or warming. Published in Russian in Fiziologiya Rastenii, 2009, Vol. 56, No. 1, pp. 35–45. The text was submitted by the authors in English.     

Physiological and antioxidant responses of <Emphasis Type="Italic">Erica multiflora</Emphasis> to drought and warming through different seasons

Climate change projections forecast a warming and an associated change in the distribution and intensity of rainfalls. In the case of the Mediterranean area, this will be reflected in more frequent and severe drought periods in the future. Within a long-term (9 years) manipulation experiment, we aimed to study the effect of the soil drought (15–20% decreased soil moisture) and warming conditions (+1°C warming) projected for the next decades onto photosynthetic rates and water relations, and onto the antioxidant and anti-stress defense capacity of Erica multiflora, a common species of the dry Mediterranean coastal scrublands, in two different seasons, spring and summer. Results indicated that none of the applied treatments was severe enough to induce a pronounced negative response of photosynthesis in this species well adapted to harsh Mediterranean conditions, but also highlighted important seasonal differences. Photosynthesis was limited by photoinhibition in spring and by stomatal closure in summer. Isoprenoid emission and the level of enzymatic and non-enzymatic antioxidants were lower in summer than in spring, whereas pigment and total phenolic contents were generally higher in summer. Volatile isoprenoid emissions were largely inhibited by drought and were not stimulated by warming. Drought and warming increased the oxidation state of ascorbate and reduced total glutathione levels in spring, but not in summer. It is concluded that E. multiflora plants can adapt to prolonged drought and warming, at least at the level simulated by our manipulative experiment, through changes in the seasonal physiological and antioxidant response of leaves.     

Warming and drought alter soil phosphatase activity and soil P availability in a Mediterranean shrubland

We conducted a field experiment simulating the warming and drought in a Mediterranean shrubland dominated by Erica multiflora and Globularia alypum with the aim to simulate the next future climate conditions predicted by the IPCC and ecophysiological models. As P is frequently a limiting nutrient in Mediterranean ecosystems, we investigated the drought and warming effects on soil phosphatases activities, soil P contents and availability, litter and leaf P concentration, and the capacity of this community to maintain soil P reserves and retain this nutrient in the ecosystem. Warming treatment increased soil and air temperature (an average of 1°C) and drought treatment decreased soil water content in one of the seasons analysed (28% in autum 2004). Warming increased (68%) the activities of soil acid phosphatases in summer and alkaline phosphatase activity (22%) in spring 2004, and increased P concentrations in E. multiflora. Instead, warming decreased P concentrations in litterfall of this same species, E. multiflora, and soil HCO₃-extractable P_i (Olsen-P_i) in some seasons, decreasing total P soil concentration (37%) after 6 years of treatment. The drought treatment did not change soil phosphatase activities, nor available P_i. The effects of climate change on soil P dynamics in Mediterranean areas will thus be strongly dependent on whether the main variable involved in the local change is warming or drought. If warming is the main change without significant changes in water availability, the increases of biological activity can accelerate plant growth, P capture by plants and increase soil-phosphatase activity, altogether decreasing P contents in soil. If drought is the main change, a reduction in P demands by plants is expected, increasing P stocks in soils.     

Novel Approaches to Study Climate Change Effects on Terrestrial Ecosystems in the Field: Drought and Passive Nighttime Warming

This article describes new approaches for manipulation of temperature and water input in the field. Nighttime warming was created by reflection of infrared radiation. Automatically operated reflective curtains covered the vegetation at night to reduce heat loss to the atmosphere. This approach mimicked the way climate change, caused by increased cloudiness and increased greenhouse gas emissions, alters the heat balance of ecosystems. Drought conditions were created by automatically covering the vegetation with transparent curtains during rain events over a 2–5-month period. The experimental approach has been evaluated at four European sites across a climate gradient. All sites were dominated (more than 50%) by shrubs of the ericaceous family. Within each site, replicated 4-m × 5-m plots were established for control, warming, and drought treatments and the effect on climate variables recorded. Results over a two-year period indicate that the warming treatment was successful in achieving an increase of the minimum temperatures by 0.4–1.2°C in the air and soil. The drought treatment resulted in a soil moisture reduction of 33%–82% at the peak of the drought. The data presented demonstrate that the approach minimizes unintended artifacts with respect to water balance, moisture conditions, and light, while causing a small but significant reduction in wind speed by the curtains. Temperature measurements demonstrated that the edge effects associated with the treatments were small. Our method provides a valuable tool for investigating the effects of climate change in remote locations with minimal artifacts.