Otília Correia

Two different strategies of Mediterranean macchia plants to avoid photoinhibitory damage by excessive radiation levels during summer drought

The adaptive strategies to high radiation and water stress of the drought tolerant evergreen sclerophylls Quercus coccifera and Arbutus unedo are compared to those of the semi-deciduous Cistus spp. (C. albidus and C. monspeliensis). Cistus spp. partially avoided drought by a marked reduction of their transpirational surface through leaf abscission during summer, when predawn water potential declined below -5.5 MPa. Chlorophyll fluorescence measurements revealed a reversible diurnal decrease of maximum photochemical efficiency of PSII (F-v/F-m), which became more accentuated during summer drought in all species. An important strategy to avoid damage by excessive radiation levels in Cistus spp. was the structural regulation of light interception through leaf angle changes, from a more horizontal orientation in spring ( 70 degrees). Horizontal orientated leaves were highly susceptible to photoinhibition, and excessive radiation often resulted in irreversible photodamage followed by leaf abscission during summer, whereas vertical leaf orientation appeared to protect the leaf from severe photoinhibition. Still, these mechanisms were not fully successful in avoiding chronic photoinhibition, and predawn F-v/F-m values remained low in Cistus spp. during summer (only exhibiting a partial overnight recovery). Evergreen sclerophylls were less susceptible to photoinhibition, and the diurnal decline in F-v/F-m remained fully reversible during drought. Structural regulation of light interception was not found to be an important strategy in these species, and only small, though significant changes in leaf angle occurred. The ecological importance of the adaptive strategies of each functional group is discussed

Characteristic patterns of chronic and dynamic photoinhibition of different functional groups in a Mediterranean ecosystem

To analyse characteristic patterns of dynamic and chronic photoinhibition within a plant community, a new technique is proposed, which is based on the long- and short-term recovery time of maximum photochemical efficiency of PSII (F v/F m) after environmental stress. Chronic photoinhibition was determined as a sustainable decrease in predawn F v/F m, occurring during periods of prolonged stress, whereas dynamic photoinhibition was assessed from the fully reversible diurnal decline in F v/F m. Applied to a Mediterranean macchia ecosystem, this definition allowed the characterization of typical annual patterns of chronic and dynamic photoinhibition. Both types of photoinhibition were highest during summer drought. However, differences emerged among the ten dominant macchia species regarding their susceptibility to chronic photoinhibition during different seasons. Chronic and dynamic photoinhibition were dependent on leaf orientation. Semi-deciduous species avoided enhanced chronic photoinhibition through a reduction of excessive light interception by vertical foliage orientation during summer, whereas evergreen sclerophylls did not exhibit pronounced structural photoprotective mechanisms. Chronic and total photoinhibition were significantly correlated with predawn and midday water potentials, respectively, and a grouping of the macchia species into three functional groups is proposed according to this relationship.

Phenological and growth patterns of the Mediterranean oak Quercus suber L.

The phenology of Quercus suber L., a dominant species of the montados in the Iberian Peninsula, was studied for 2 years in southwest Portugal. The seasonal progression of phenological events was analyzed in seven trees. Selected branches were examined monthly for shoot elongation, leaf number, branching, flowering, and fruiting. Radial stem growth and specific leaf weight were also studied. Active growth was observed from early spring to early summer. Reserves accumulated during winter and high photosynthetic activity in early spring apparently supported this strong development. The growth flush started with stem radial increment, which seemed to be impaired by spring rainfall. Male inflorescence production was the next phenological event. Old leaves were shed during new twig and leaf emergence. Shoot elongation and the number of new leaves produced were well correlated with the previous-year shoots length, and were not clearly related to climatic factors. Radial growth resumed in autumn at a lower rate than in the previous spring, a possible consequence of a reserve depletion due to lower photosynthetic production in summer and investment on fruit maturation, which was complete by late autumn. Premature and excessive new leaf production were apparently subjected to self-pruning strategies related to the development of each trees crown. Younger cork-oaks produced shorter and fewer shoots per module, and more sclerophyllous leaves than the older ones. A high intra-specific variability was observed in all the results.

Water relations of cork-oak (Quercus suber L.) under natural conditions

Daily and annual courses of leaf transpiration, stomatal conductance and shoot water potential of four Quercus suber individuals were compared in a semi-natural stand in southwest Portugal, from spring 1989 to early summer 1990. The trees investigated showed annual patterns typical of evergreen sclerophyllous species but varied in their range of stomatal operation. This appeared to be related to differences in hydraulic conductivity in the root-to-leaf pathway. Maximum stomatal conductance and transpiration rates occurred from March to June. Water stress was found to be moderate and winter cold stress due to low air and soil temperatures appeared to have an influence on plant water balance through their effects on flow resistances.

Nutrient dynamics in crown tissues of cork-oak (Quercus suber L.)

The seasonal fluctuation of N, P, K, Ca and Mg in leaves, young stems and fruits of young and matureQuercus suber trees growing at the same site was studied. Nutrient dynamics within the tree appeared to be linked to phenological events. Leaves maximized their nutrient contents throughout summer whereas stems maintained rather constant values during most of the year. Before the spring growth flush they apparently acted as temporary reservoirs for some nutrients, translocated from leaves or other organs, so that such nutrients would be easily invested in the new tissues. A significant investment was made in sink organs like flowers and fruits. The tree probably relies not only on leaves but also on other nutrient reservoirs like older branches and roots for an adequate supply to those sinks and to the spring growth. The importance of the sampling dates and the basis of expression used for nutrient concentrations has also been emphasized. Nutrient contents and seasonal trends of the studied trees were similar to those of other evergreen oaks, although somewhat higher values for N, K, and Mg were found. Moreover, soil nutrient contents in the studied “montado” were much lower than in other Iberic oak stands. However the nutrient status and requirements of cork-oak suggest high (or at least adequate) nutrient availability to the trees. The mechanisms by which this could be achieved are discussed.

Photoinhibition in cork-oak leaves under stress: influence of the bark-stripping on the chlorophyll fluorescence emission inQuercus suber L.

Quercus suber is the primary source for industrial cork and becomes bark-stripped every 9–10 years. Recurring cork extraction is a major stress factor and the large water loss from the stripped trunk surface may affect the water balance and tree productivity. To evaluate the effect of bark-stripping, fluorescence emission and stomatal conductance of leaves were determined in groups of bark-stripped and control trees. Fv/Fm ratio was found to be significantly lower in bark-stripped trees indicating a reduced photosynthetic efficiency of PSII. Photosynthesis was not found to be stomata limited. The reduction in Fv/Fm resulted from a decline in maximum and variable fluorescence while the initial fluorescence of the dark-adapted state (Fo) remained constant. A general decline in photosynthetic efficiency of PSII was found in all trees during the summer, probably reflecting the prolonged environmental stresses during a hot and dry season. Additional stress caused by the bark-stripping seems to enhance the susceptibility to photoinhibition of the trees.

Lichens as ecological indicators in urban areas: beyond the effects of pollutants

Summary 1. In this work, we hypothesized that in urban areas with reduced pollution levels, local climatic conditions can be an important driver of lichen diversity. Thus, lichen functional groups could be used to study the effects of the urban heat island, characterized by higher temperature and lower humidity in urban than in peri-urban areas. To test this hypothesis, we sampled functional groups of epiphytic lichens in 48 forest remnants of a Mediterranean urban area (Almada, Portugal). 2. Of all tested functional groups (including those most closely associated with air pollution), groups related to water requirements showed the most significant correlations with surrounding artificial areas, roads and forest. These results suggest that the current major driver of lichen diversity is the climate within the urban area rather than air pollution, as frequently assumed. 3. The analysis of local climate using meteorological data of temperature and humidity confirmed the existence of urban heat island in the study area. 4. Synthesis and applications. The analysis of lichen functional groups gave an integrated response to the climatic modifications occurring in urban areas, namely to the urban heat island, suggesting that lichens can be used as a tool to evaluate the impact of urban areas on local climate and the effectiveness of mitigation and adaptation strategies if pollution levels are low.

Sexual dimorphism, sex ratio and spatial distribution of male and female shrubs in the dioecious speciesPistacia lentiscus L.

The population sex ratio, spatial distribution, relative growth rate, canopy size and morphology, and nitrogen content of leaves of male and female individuals were compared in seven populations of the dioecious shrubPistacia lentiscus L. occurring in different localities of the Serra da Arrábida Natural Park, Portugal. The expected pattern of male preponderance in stressful habitats was tested.Less perturbed areas, with a well developed vegetation cover, had male-biased sex ratios, while in former agricultural areas there were no significant differences between the number of male and female plants. Our results failed to reveal any significant differences in the morphological variables between sexes except in the number of basal stems. Male plants also had a significantly higher chlorophyll content (chl a+b) than female plants. But when plants were sorted into large and small individuals, larger plants showed a significantly higher leaf area index and a significantly higher number of basal stems. There were no significant differences in the leaf nitrogen content between male and female plants from June to August, but such differences appeared (P<0.001) in September and October corresponding to the production of mature fruits.Differences in sex-ratio may be explained by different resource allocation patterns of the two sexes, which favoured male plants in the less perturbed environments. In agricultural areas farmers could also have favoured female plants in former times.

Evaluating green infrastructure in urban environments using a multi-taxa and functional diversity approach.

Forested areas within cities host a large number of species, responsible for many ecosystem services in urban areas. The biodiversity in these areas is influenced by human disturbances such as atmospheric pollution and urban heat island effect. To ameliorate the effects of these factors, an increase in urban green areas is often considered sufficient. However, this approach assumes that all types of green cover have the same importance for species. Our aim was to show that not all forested green areas are equal in importance for species, but that based on a multi-taxa and functional diversity approach it is possible to value green infrastructure in urban environments. After evaluating the diversity of lichens, butterflies and other-arthropods, birds and mammals in 31 Mediterranean urban forests in south-west Europe (Almada, Portugal), bird and lichen functional groups responsive to urbanization were found. A community shift (tolerant species replacing sensitive ones) along the urbanization gradient was found, and this must be considered when using these groups as indicators of the effect of urbanization. Bird and lichen functional groups were then analyzed together with the characteristics of the forests and their surroundings. Our results showed that, contrary to previous assumptions, vegetation density and more importantly the amount of urban areas around the forest (matrix), are more important for biodiversity than forest quantity alone. This indicated that not all types of forested green areas have the same importance for biodiversity. An index of forest functional diversity was then calculated for all sampled forests of the area. This could help decision-makers to improve the management of urban green infrastructures with the goal of increasing functionality and ultimately ecosystem services in urban areas.

Optimization of Landscape Services under Uncoordinated Management by Multiple Landowners

Landscapes are often patchworks of private properties, where composition and configuration patterns result from cumulative effects of the actions of multiple landowners. Securing the delivery of services in such multi-ownership landscapes is challenging, because it is difficult to assure tight compliance to spatially explicit management rules at the level of individual properties, which may hinder the conservation of critical landscape features. To deal with these constraints, a multi-objective simulation-optimization procedure was developed to select non-spatial management regimes that best meet landscape-level objectives, while accounting for uncoordinated and uncertain response of individual landowners to management rules. Optimization approximates the non-dominated Pareto frontier, combining a multi-objective genetic algorithm and a simulator that forecasts trends in landscape pattern as a function of management rules implemented annually by individual landowners. The procedure was demonstrated with a case study for the optimum scheduling of fuel treatments in cork oak forest landscapes, involving six objectives related to reducing management costs (1), reducing fire risk (3), and protecting biodiversity associated with mid- and late-successional understories (2). There was a trade-off between cost, fire risk and biodiversity objectives, that could be minimized by selecting management regimes involving ca. 60% of landowners clearing the understory at short intervals (around 5 years), and the remaining managing at long intervals (ca. 75 years) or not managing. The optimal management regimes produces a mosaic landscape dominated by stands with herbaceous and low shrub understories, but also with a satisfactory representation of old understories, that was favorable in terms of both fire risk and biodiversity. The simulation-optimization procedure presented can be extended to incorporate a wide range of landscape dynamic processes, management rules and quantifiable objectives. It may thus be adapted to other socio-ecological systems, particularly where specific patterns of landscape heterogeneity are to be maintained despite imperfect management by multiple landowners.