Alberto Vilagrosa

Pines and oaks in the restoration of Mediterranean landscapes of Spain: New perspectives for an old practice — a review

Pines have been extensively used for land restoration in the Mediterranean basin and in other parts of the world, since the late 19th century. The theoretical basis supporting pine utilisation was its stress-tolerant and pioneer features, and their attributed role of facilitating the development of late-successional hardwoods in the long-term. In the present work, the use of pines and hardwoods in forest restoration is discussed in the frame of the current disturbance regime and social demands for Mediterranean forests. Large pine plantations have recently disappeared because of their sensitivity to fire (e.g., Pinus nigra) or because of the short fire-intervals (e.g., Pinus halepensis). Combined pine and oak plantations are proposed for degraded land restoration on the basis of the complementary features of both groups of species. Seeding and containerised seedling plantation, soil amendments and plantation techniques to reduce transplant shock are evaluated for reforestation under water-stressing conditions, on the basis of several experiments performed in eastern Spain. Both P. halepensis and Quercus ilex are tested.

Morphological traits and water use strategies in seedlings of Mediterranean coexisting species

The distribution of plants is associated with their different patterns of response to their environment. Mediterranean plants have evolved a number of morphological and physiological adaptations that determine their ability to survive and grow, being an effective water uptake and use important factors for drought resistance. In this article, we evaluated interspecific differences in morphology, biomass allocation, and architectural traits and their relationship with water use strategies in seedlings of seven co-occurring Mediterranean species (Anthyllis cytisoides L., Genista scorpius L. DC., Myrtus communis L., Pistacia lentiscus L., Rosmarinus officinalis L., Spartium junceum L. and Ulex parviflorus Pourr.). The results showed that morphological root features vary among species and they are significantly correlated with root hydraulic conductance and leaf gas exchange variables. Species with high specific root length (SRL) showed a low hydraulic conductance per root length (KRRL) but high specific hydraulic conductance (KAs). M. communis and P. lentiscus showed the most contrasting water use patterns with respect to the other species studied. The results are not affected when considering phylogenetic relatedness. Thus, the variability observed in root hydraulic properties and leaf gas exchange suggests important mechanisms for understanding species coexistence in water-limited ecosystems.

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.

Perspectives in dryland restoration: approaches for climate change adaptation

Reforestation efforts in dryland ecosystems frequently encounter drought and limited soil productivity, although both factors usually interact synergistically to worsen water stress for outplanted seedlings. Land degradation in drylands (e.g. desertification) usually reduces soil productivity and, especially, soil water availability. In dry sub-humid regions, forest fires constitute a major disturbance affecting ecosystem dynamics and reforestation planning. Climate change projections indicate an increase of drought and more severe fire regime in many dryland regions of the world. In this context, the main target of plantation technology development is to overcome transplant shock and likely adverse periods, and in drylands this is mostly related to water limitations. In this paper, we discuss some selected steps that we consider critical for improving success in outplanting woody plants, both under current and projected climate change conditions including: (1) Plant species selection, (2) Improved nursery techniques, and (3) Improved planting techniques. The number of plant species used in reforestation is increasing rapidly, moving from a reduced set of well-known, easy-to-grow, widely used species, to a large variety of promising native species. Available technologies allow for reintroducing native plants and recovering critical ecosystem functions for many degraded drylands. However, climate change projections introduce large uncertainties about the sustainability of current reforestation practices. To cope with these uncertainties, adaptive restoration approaches are suggested, on the basis of improved plant quality, improved techniques for optimizing rain use efficiency in plantations, and exploring native plant species, including provenances and genotypes, for their resilience to fire and water use efficiency.

Towards understanding resprouting at the global scale.

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

Physiological differences explain the co-existence of different regeneration strategies in Mediterranean ecosystems

• Fire and drought are selective driving forces in Mediterranean plants, and thus their ability to resprout or recruit after these disturbances is of paramount importance. The contrast in regeneration niche between resprouters and seeders, and each groups different root characteristics, suggest that they are subjected to different degrees of environmental stress and, consequently, to different evolutionary forces. • We compared leaf traits, xylem traits related to hydraulic efficiency and vulnerability to cavitation, and the physiological response to an imposed drought between seedlings of resprouters and seedlings of seeders. We used 12 species co-existing in Mediterranean basin ecosystems. • Major differences were found in the xylem architecture and leaf traits, and in the response to drought conditions. Seeders were more efficient at transporting water to leaves but formed, in turn, a safer xylem network. They also presented higher photosynthesis and transpiration rates, and earlier stomatal closure with drought, but good leaf dehydration tolerance. • Seeders and resprouters can be considered two syndromes whose different functional characteristics are related to water availability and drought responses. These characteristics, together with their differences in rooting habits, account for their distinct regeneration niches and, thus, their co-existence.

Leaf physiological traits in relation to resprouter ability in the Mediterranean Basin

In Mediterranean ecosystems, fire is a strong selective agent among plants, and the different post-fire regeneration strategies (e.g. resprouting and non-resprouting) have implications for other plant traits. Because young plants of non-resprouters need to grow quickly and mature well before the next fire, we predict that they should possess leaf traits related to increased efficiency in growth and resource acquisition compared with resprouter species. To test this hypothesis, we measured specific leaf area, leaf nitrogen and carbon concentrations and leaf physiological traits, including gas exchange parameters and chlorophyll fluorescence, in 19 Mediterranean species cultivated in a common garden. Both cross-species and phylogenetically informed analyses suggest that non-resprouters have better physiological performance at the leaf level (i.e. higher photosynthetic capacity) than resprouters. All these results suggest that non-resprouter species are able to take greater advantage for vegetative growth and carbon fixation than resprouters during periods when water is readily available. The contrasted physiological differences between resprouters and non-resprouters reinforce the idea that these two syndromes are functionally different (i.e. they are functional types).

The role of nutrients for improving seedling quality in drylands

AbstractForest plantations have been extensively used to combat desertification. In drylands, harsh climate conditions and unfertile soils often preclude seedling establishment. The improvement in seedling quality by manipulating nutrient availability could contribute to increase planting success. However, morpho-functional traits defining optimum seedling quality in drylands, and the fertilization schemes to achieve them, are still under discussion. Several studies suggest that well fertilized seedlings may perform better than nutrient limited seedlings in these environments. However, recent works have shown opposite results. In this review, we discuss the concept of seedling quality in drylands based on an evaluation of the effects of nutrient manipulation on seedling morpho-functional traits and field performance. According to existing data, we hypothesize that nutrient-limited small seedlings may be better adapted to arid environments and unfavorable microsites, where access to water is uncertain and a conservative water use strategy may be advantageous. In contrast, in dry sub-humid areas, areas with deep soils, protected from excess radiation, and areas where irrigation is feasible, well-fertilized big seedlings with high root growth potential may have more chances of success. We discuss this theory in the context of the multiple objectives of dryland restoration and the environmental constrains posed by these areas, and identify knowledge gaps that should be targeted to test our hypothesis.

Xylem Cavitation and Embolism in Plants Living in Water-Limited Ecosystems

Water deficit is considered the main limiting factor for the establishment, survival, and growth of plants mainly in water-limited ecosystems. Plants have evolved a wide range of morphologic and functional mechanisms to adapt to arid environments. However, if the tension in the xylem conduits becomes too high, thus xylem cavitation can occur i.e., water column breakage. This results in the hydraulic disconnection of leaves and above-ground parts from roots because xylem conduits are filled with air and water vapor, and this phenomenon is called embolism. Therefore, the resistance of the xylem to cavitation and embolism is of paramount importance for plant functioning. In this chapter, we will review the role of plant hydraulics and xylem cavitation in the context of water-limited ecosystems and their relationship with other plant functional traits and with survival capacity. These topics will be analyzed and discussed on the basis of current knowledge and our research experiences.

Root architecture and hydraulic conductance in nutrient deprived Pistacia lentiscus L. seedlings

Plants respond to low nutrient availability by modifying root morphology and root system topology. Root responses to nitrogen (N) and phosphorus (P) limitation may affect plant capacity to withstand water stress. But studies on the effect of nutrient availability on plant ability to uptake and transport water are scarce. In this study, we assess the effect of nitrogen and phosphorus limitation on root morphology and root system topology in Pistacia lentiscus L seedlings, a common Mediterranean shrub, and relate these changes to hydraulic conductivity of the whole root system. Nitrogen and phosphorus deprivation had no effect on root biomass, but root systems were more branched in nutrient limited seedlings. Total root length was higher in seedlings subjected to phosphorus deprivation. Root hydraulic conductance decreased in nutrient-deprived seedlings, and was related to the number of root junctions but not to other architectural traits. Our study shows that changes in nutrient availability affect seedling water use by modifying root architecture. Changes in nutrient availability should be taken into account when evaluating seedling response to drought.