Question: A set of easily-measured ('soft') plant traits has been identified as potentially useful predictors of ecosystem functioning in previous studies. Here we aimed to discover whether the screening techniques remain operational in widely contrasted circumstances, to test for the existence of axes of variation in the particular sets of traits, and to test for their links with 'harder' traits of proven importance to ecosystem functioning. Location: central-western Argentina, central England, northern upland Iran, and northeastern Spain. Recurrent patterns of ecological specialization: Through ordination of a matrix of 640 vascular plant taxa by 12 standardized traits, we detected similar patterns of specialization in the four floras. The first PCA axis was identified as an axis of resource capture, usage and release. PCA axis 2 appeared to be a size-related axis. Individual PCA for each country showed that the same traits remained valuable as predictors of resource capture and utilization in all of them, despite their major differences in climate, biogeography and land-use. The results were not significantly driven by particular taxa: the main traits determining PCA axis 1 were very similar in eudicotyledons and monocotyledons and Asteraceae, Fabaceae and Poaceae. Links between recurrent suites of 'soft' traits and 'hard' traits: The validity of PCA axis 1 as a key predictor of resource capture and utilization was tested by comparisons between this axis and values of more rigorously established predictors ('hard' traits) for the floras of Argentina and England. PCA axis 1 was correlated with variation in relative growth rate, leaf nitrogen content, and litter decomposition rate. It also coincided with palatability to model generalist herbivores. Therefore, location on PCA axis 1 can be linked to major ecosystem processes in those habitats where the plants are dominant. Conclusion: We confirm the existence at the global scale of a major axis of evolutionary specialization, previously recog-nised in several local floras. This axis reflects a fundamental trade-off between rapid acquisition of resources and conservation of resources within well-protected tissues. These major trends of specialization were maintained across different environmental situations (including differences in the proximate causes of low productivity, i.e. drought or mineral nutrient deficiency). The trends were also consistent across floras and major phylogenetic groups, and were linked with traits directly relevant to ecosystem processes.

In this study we have addressed the effect of nitrogen fertilisation and shading used during nursery growth on the functional quality of Quercus ilex L. (holm oak) seedlings and their outplanting performance in an abandoned continental Mediterranean cropland. Three N fertilisation regimes: high and low fertilisation (154 and 34 mg N per plant, respectively) and no fertilisation, combined with two shade levels (full sun and 45% shade) were studied. Fertilisation increased the shoot size, shoot to root mass ratio (S/R), root N, and both shoot and root P concentration of the plants produced in the nursery. Fertilisation also enhanced the capacity of new root formation. Neither tissue K nor root mass was affected by N fertilisation. Shading increased S/R and root K concentration but reduced both shoot and root N concentration. Two years after planting, unfertilised seedlings had higher mortality and grew significantly less than those cultivated at both a low and a high fertilisation level. Shading during nursery growth had no influence on the outplanting performance of holm oak seedlings. Plants with a larger shoot and higher S/R had lower mortality and grew faster than those with a smaller shoot and lower S/R. Outplanting performance of holm oak seedlings was related with root N and P concentration and with RGC. No relationship was found with either shoot nutrient concentration or root mass. We conclude that high N fertilisation of holm oak seedlings in the nursery can improve their early field establishment in the afforestation Mediterranean abandoned croplands. #

Reduction in plant size and tissue nutrient concentration is widely considered to increase 30 seedling drought resistance in dry and oligotrophic plantation sites. However, much evidence 31 indicates that increase in size and tissue nutrient concentration improves seedling survival in 32 Mediterranean forest plantations. This suggests that the ecophysiological processes and 33 functional attributes relevant for early seedling survival in Mediterranean climate must be 34 reconsidered. We propose a physiological conceptual model for seedling survival in 35 Mediterranean-climate plantations to provide a physiological explanation of the frequent 36 positive relationship between outplanting performance and seedling size and nutrient 37 concentration. The model considers the physiological processes outlined in the plantation 38 establishment model of Burdett (1990), but incorporates other physiological processes that 39 drive seedling survival, such as N remobilization, carbohydrate storage and plant hydraulics.40 The model considers that seedling survival in Mediterranean climates is linked to high growth 41 capacity during the wet season. The model is for container plants and is based on three main 42 principles, 1) Mediterranean climates are not dry the entire year but usually have two 43 seasons of contrasting water availability; 2) summer drought is the main cause of seedling 44 mortality; in this context deep and large roots is a key trait for avoiding lethal water stress; 3) 45 attainment of large root systems in the dry season is promoted when seedlings have high 46 growth during the wet season. High growth is achieved when seedlings can divert large 47 amount of resources to support new root and shoot growth. Functional traits that confer high 48 photosynthesis, nutrient remobilization capacity, and non-structural carbohydrate storage 49 promote high growth. Increases in seedling size and nutrient concentration, strongly affect 50 these physiological processes. Traits that confer high drought resistance are of low value 51 during the wet season because hinder growth capacity. We provide specific evidence to 52 support the model and finally we discuss its implications and the factors that may alter the 53 frequent increase in performance with increase in seedling size and tissue nutrient 54 concentration.

Summary 1.Competitive and facilitative interactions shape plant communities. Whereas a number of studies have addressed competition and direct facilitation among plants in dry ecosystems, indirect facilitation has received little attention. 2. We investigated the relative importance of direct and indirect facilitation by the nurse plant Retama sphaerocarpa on late-successional Quercus ilex seedlings mediated by herb suppression in a Mediterranean shrubland in 2006 and 2007. We also studied whether facilitation outcome depended on the size of the facilitated seedlings. 3. A field experiment was carried out to test the effect of (i) position of Q. ilex seedling with respect to shrub canopy (under shrubs or in gaps), (ii) herb competition (presence or absence), and (iii) seedling size. 2006 was an average rainfall year while 2007 had a much more humid spring and a dryer summer than 2006. 4. In both years, nurse shrubs reduced seedling mortality whereas herbs increased it. In the average rainfall year, seedling mortality under shrubs was unaffected by herbs whereas in gaps it was significantly higher in presence of herbs. This showed that the nurse shrub indirectly facilitated the seedlings by reducing the competitive capacity of herbs. Conversely, facilitation was predominately direct during the humid spring and dry summer year since herbs hindered seedling survival similarly under the nurse shrub and in gaps. The nurse shrub directly facilitated the seedlings by reducing seedling photoinhibition and water stress. 5. Improvement of environmental conditions by Retama benefited smaller seedlings but not larger seedlings since the nurse shrub reduced mortality of smaller seedlings relative to that in gaps, but this effect was not observed for larger seedlings. This indicates that individuals within a seedling population may not have the same response to facilitation. 6. Synthesis. Both indirect and direct facilitation are important mechanisms for Q. ilex regeneration in Retama shrubland and their importance seems to vary with climatic conditions. Indirect facilitation by release of herb competition under nurse shrubs is important in years of dry springs when competition between nurse shrubs and herbs is high, whereas direct facilitation mediated by microclimate amelioration increases with summer aridity.

AbstractmStem xylem features in two evergreen Quercus species (Q. coccifera and Q. ilex) and a deciduous one (Q. faginea) were analysed along an Atlantic-Mediterranean climatic gradient in which rainfall and winter cold experience strong variation. Mean maximum vessel diameter, vessel density, vessel element length, xylem transverse sectional area, Huber value (xylem transverse sectional area per leaf area unit), theoretical leaf specific conductivity (estimated hydraulic conductance per leaf area unit) and total leaf area were determined in 3-year-old branches. Q. faginea presented the widest vessels and the highest theoretical leaf specific conductivity while Q. coccifera showed the lowest total leaf area and the highest Huber value. Studied features did not exhibit significant correlations with mean minimum January temperature in any species but did show significant relationships with rainfall. In Q. coccifera, mean maximum vessel diameter, vessel element length and theoretical leaf specific conductivity increased with higher rainfall while vessel density decreased. Mean maximum vessel diameter and total leaf area in Q. ilex increased with precipitation whereas variables of Q. faginea did not show any significant trend. Results suggest that aridity, rather than minimum winter temperature, controls stem xylem responses in the studied evergreen species. Q. faginea traits did not show any response to precipitation, probably because this species develops deep roots, which in turn makes edaphic and topographic factors more important in the control of soil water availability. In response to aridity Q. coccifera only exhibits adjustment at a xylem level by reducing its water transport capacity through a reduction of vessel diameter without changing the amount of xylem tissue or foliage, whereas Q. ilex adjusts its water transport capacity in parallel to the foliage area.