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Models derived from satellite image data are needed to monitor the status of terrestrial ecosystems across large spatial scales. However, a remote sensing-based approach to quantify soil multifunctionality at the global scale is missing despite significant research efforts on this topic. A major constraint for doing so is the availability of suitable global-scale field data to calibrate remote sensing indicators (RSI) and, to a lesser extent, the sensitivity of spectral data of available satellite sensors to soil background and atmospheric conditions. Here, we aimed to develop a soil multifunctionality model to monitor global drylands coupling ground data on 14 soil functions of 222 dryland areas from six continents to 18 RSI derived from a time series (2006–2013) Landsat dataset. Among the RSI evaluated, the chlorophyll absorption ratio index was the best predictor of soil multifunctionality in single-variable-based models (r = 0.66, P < 0.01, NMRSE = 0.17). However, a multi-variable RSI model combining the chlorophyll absorption ratio index, the global environment monitoring index and the canopy-air temperature difference improved the accuracy of quantifying soil multifunctionality (r = 0.73, P < 0.01, NMRSE = 0.15). Furthermore, the correlation between RSI and soil variables shows a wide range of accuracy with upper and lower values obtained for AMI (r = 0.889, NMRSE = 0.05) and BGL (r = 0.685, NMRSE = 0.18) respectively. Our results provide new insights on assessing soil multifunctionality using RSI that may help to monitor temporal changes in the functioning of global drylands effectively.

Previous research indicates that some important cocoa cultivated areas in West Africa will become unsuitable for growing cocoa in the next decades. However, it is not clear if this change will be mirrored by the shade tree species that could be used in cocoa-based agroforestry systems (C-AFS). We characterized current and future patterns of habitat suitability for 38 tree species (including cocoa), using a consensus method for species distribution modelling considering for the first time climatic and soil variables. The models projected an increase of up to 6% of the potential suitable area for cocoa by 2060 compared to its current suitable area in West Africa. Furthermore, the suitable area was highly reduced (14.5%) once considering only available land-use not contributing to deforestation. Regarding shade trees, 50% of the 37 shade tree species modelled will experience a decrease in geographic rate extent by 2040 in West Africa, and 60% by 2060. Hotspots of shade tree species richness overlap the current core cocoa production areas in Ghana and Côte d’Ivoire, suggesting a potential mismatch for the outer areas in West Africa. Our results highlight the importance of transforming cocoa-based agroforestry systems by changing shade tree species composition to adapt this production systems for future climate conditions.

1. Multiple ecosystem functions need to be considered simultaneously to manage and protect the several ecosystem services that are essential to people and their environments. Despite this, cost effective, tangible, relatively simple and globally relevant methodologies to monitor in situ soil multifunctionality, that is, the provision of multiple ecosystem functions by soils, have not been tested at the global scale. 2. We combined correlation analysis and structural equation modelling to explore whether we could find easily measured, field‐based indicators of soil multifunctionality (measured using functions linked to the cycling and storage of soil carbon, nitrogen and phosphorus). To do this, we gathered soil data from 120 dryland ecosystems from five continents. 3. Two soil surface attributes measured in situ (litter incorporation and surface aggregate stability) were the most strongly associated with soil multifunctionality, even after accounting for geographic location and other drivers such as climate, woody cover, soil pH and soil electric conductivity. The positive relationships between surface stability and litter incorporation on soil multifunctionality were greater beneath the canopy of perennial vegetation than in adjacent, open areas devoid of vascular plants. The positive associations between surface aggregate stability and soil functions increased with increasing mean annual temperature. 4. Synthesis and applications. Our findings demonstrate that a reduced suite of easily measured in situ soil surface attributes can be used as potential indicators of soil multifunctionality in drylands world‐wide. These attributes, which relate to plant litter (origin, incorporation, cover), and surface stability, are relatively cheap and easy to assess with minimal training, allowing operators to sample many sites across widely varying climatic areas and soil types. The correlations of these variables are comparable to the influence of climate or soil, and would all

1. Multiple ecosystem functions need to be considered simultaneously to manage and protect the several ecosystem services that are essential to people and their environments. Despite this, cost effective, tangible, relatively simple and globally relevant methodologies to monitor in situ soil multifunctionality, that is, the provision of multiple ecosystem functions by soils, have not been tested at the global scale. 2. We combined correlation analysis and structural equation modelling to explore whether we could find easily measured, field‐based indicators of soil multifunctionality (measured using functions linked to the cycling and storage of soil carbon, nitrogen and phosphorus). To do this, we gathered soil data from 120 dryland ecosystems from five continents. 3. Two soil surface attributes measured in situ (litter incorporation and surface aggregate stability) were the most strongly associated with soil multifunctionality, even after accounting for geographic location and other drivers such as climate, woody cover, soil pH and soil electric conductivity. The positive relationships between surface stability and litter incorporation on soil multifunctionality were greater beneath the canopy of perennial vegetation than in adjacent, open areas devoid of vascular plants. The positive associations between surface aggregate stability and soil functions increased with increasing mean annual temperature. 4. Synthesis and applications. Our findings demonstrate that a reduced suite of easily measured in situ soil surface attributes can be used as potential indicators of soil multifunctionality in drylands world‐wide. These attributes, which relate to plant litter (origin, incorporation, cover), and surface stability, are relatively cheap and easy to assess with minimal training, allowing operators to sample many sites across widely varying climatic areas and soil types. The correlations of these variables are comparable to the influence of climate or soil, and would all

Key message This study reinforces the existence of the leaf economics spectrum in Mediterranean woody species, and demonstrates the strong influence of phylogeny, leaf habit and environmental context as main drivers of variability in structural and nutrient traits of leaves. Abstract Leaf structural and nutrient traits are key attributes of plant ecological strategies, as these traits are related to resource-use strategies and plant growth. However, leaf structure and nutrient composition can vary among different habitats, leaf habits or phylogenetic groups. In this study, we measured 13 leaf traits (one structural—leaf mass per area, LMA—and 12 nutrient traits) in 98 Mediterranean woody species growing over a wide range of environmental conditions, with the final aim of discerning the main causes of leaf trait variability. The variance decomposition results show that phylogeny, leaf habit and habitat type affected in several ways the structural and nutrient traits studied. Leaf nutrient concentrations are strongly positively correlated amongst themselves, and negatively correlated with LMA, in accordance with the “leaf economics spectrum”. We found that leaf habit and phylogeny were important causes of variation in LMA and in a broad number of leaf nutrients (i.e., C, N, Mg, S, K), while other micronutrients seemed to be more dependent on the environment (i.e., Cu and Mn). In summary, our study reinforces the existence of the leaf economics spectrum in a broad pool of Mediterranean woody species, and demonstrates the strong influence of phylogeny, leaf habit and environmental context as the main drivers of variability in some leaf structural and nutrient traits.

Intransitive competition networks, those in which there is no single best competitor, may ensure species coexistence. However, their frequency and importance in maintaining diversity in real-world ecosystems remain unclear. We used two large data sets from drylands and agricultural grasslands to assess: (1) the generality of intransitive competition, (2) intransitivity–richness relationships and (3) effects of two major drivers of biodiversity loss (aridity and land-use intensification) on intransitivity and species richness. Intransitive competition occurred in > 65% of sites and was associated with higher species richness. Intransitivity increased with aridity, partly buffering its negative effects on diversity, but was decreased by intensive land use, enhancing its negative effects on diversity. These contrasting responses likely arise because intransitivity is promoted by temporal heterogeneity, which is enhanced by aridity but may decline with land-use intensity. We show that intransitivity is widespread in nature and increases diversity, but it can be lost with environmental homogenisation.

Aim The global spread of woody plants into grasslands is predicted to increase over the coming century. While there is general agreement regarding the anthropogenic causes of this phenomenon, its ecological consequences are less certain. We analysed how woody vegetation of differing cover affects plant diversity (richness and evenness) and the surrogates of multiple ecosystem processes (multifunctionality) in global drylands, and how these change with aridity. Location Two hundred and twenty-four dryland sites from all continents except Antarctica, widely differing in their environmental conditions (from arid to dry-subhumid sites) and relative woody cover (from 0 to 100%). Methods Using a standardized field survey, we measured the cover, richness and evenness of perennial vegetation. At each site, we measured 14 soil variables related to fertility and the build-up of nutrient pools. These variables are critical for maintaining ecosystem functioning in drylands. Results Species richness and ecosystem multifunctionality were strongly related to woody vegetation, with both variables peaking at a relative woody cover (RWC) of 41–60%. This relationship shifted with aridity. We observed linear positive effects of RWC in dry-subhumid sites. These positive trends shifted to hump-shaped RWC–diversity and multifunctionality relationships under semi-arid environments. Finally, hump-shaped (richness, evenness) or linear negative (multifunctionality) effects of RWC were found under the most arid conditions. Main conclusions Plant diversity and multifunctionality peaked at intermediate levels of woody cover, although this relationship became increasingly positive in wetter environments. This comprehensive study accounts for multiple ecosystem attributes across a range of levels of woody cover and environmental conditions. Our results help us to reconcile contrasting views of woody encroachment found in the current literature and can be used to improve predictions of the likely e

Plant–plant interactions are driven by environmental conditions, evolutionary relationships (ER) and the functional traits of the plants involved. However, studies addressing the relative importance of these drivers are rare, but crucial to improve our predictions of the effects of plant–plant interactions on plant communities and of how they respond to differing environmental conditions. To analyze the relative importance of – and interrelationships among – these factors as drivers of plant–plant interactions, we analyzed perennial plant co-occurrence at 106 dryland plant communities established across rainfall gradients in nine countries. We used structural equation modelling to disentangle the relationships between environmental conditions (aridity and soil fertility), functional traits extracted from the literature, and ER, and to assess their relative importance as drivers of the 929 pairwise plant–plant co-occurrence levels measured. Functional traits, specifically facilitated plants’ height and nurse growth form, were of primary importance, and modulated the effect of the environment and ER on plant–plant interactions. Environmental conditions and ER were important mainly for those interactions involving woody and graminoid nurses, respectively. The relative importance of different plant–plant interaction drivers (ER, functional traits, and the environment) varied depending on the region considered, illustrating the difficulty of predicting the outcome of plant–plant interactions at broader spatial scales. In our global-scale study on drylands, plant–plant interactions were more strongly related to functional traits of the species involved than to the environmental variables considered. Thus, moving to a trait-based facilitation/competition approach help to predict that: (1) positive plant–plant interactions are more likely to occur for taller facilitated species in drylands, and (2) plant–plant interactions within woody-dominated ecosystems might be more sen

The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems1. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes1,2,3,4,5. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability6,7,8. The increase in aridity predicted for the twenty-first century in many drylands worldwide9,10,11 may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients12,13,14. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition12,13,14. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

Dryland ecosystems account for ca. 27% of global soil organic carbon (C) reserves, yet it is largely unknown how climate change will impact C cycling and storage in these areas. In drylands, soil C concentrates at the surface, making it particularly sensitive to the activity of organisms inhabiting the soil uppermost levels, such as communities dominated by lichens, mosses, bacteria and fungi (biocrusts). We conducted a full factorial warming and rainfall exclusion experiment at two semiarid sites in Spain to show how an average increase of air temperature of 2–3 °C promoted a drastic reduction in biocrust cover (ca. 44% in 4 years). Warming significantly increased soil CO2 efflux, and reduced soil net CO2 uptake, in biocrust-dominated microsites. Losses of biocrust cover with warming through time were paralleled by increases in recalcitrant C sources, such as aromatic compounds, and in the abundance of fungi relative to bacteria. The dramatic reduction in biocrust cover with warming will lessen the capacity of drylands to sequester atmospheric CO2. This decrease may act synergistically with other warming-induced effects, such as the increase in soil CO2 efflux and the changes in microbial communities to alter C cycling in drylands, and to reduce soil C stocks in the mid to long term.

While much is known about the factors that control each component of the terrestrial nitrogen (N) cycle, it is less clear how these factors affect total N availability, the sum of organic and inorganic forms potentially available to microorganisms and plants. This is particularly true for N-poor ecosystems such as drylands, which are highly sensitive to climate change and desertification processes that can lead to the loss of soil nutrients such as N. We evaluated how different climatic, abiotic, plant and nutrient related factors correlate with N availability in semiarid Stipa tenacissima grasslands along a broad aridity gradient from Spain to Tunisia. Aridity had the strongest relationship with N availability, suggesting the importance of abiotic controls on the N cycle in drylands. Aridity appeared to modulate the effects of pH, plant cover and organic C (OC) on N availability. Our results suggest that N transformation rates, which are largely driven by variations in soil moisture, are not the direct drivers of N availability in the studied grasslands. Rather, the strong relationship between aridity and N availability could be driven by indirect effects that operate over long time scales (decades to millennia), including both biotic (e.g. plant cover) and abiotic (e.g. soil OC and pH). If these factors are in fact more important than short-term effects of precipitation on N transformation rates, then we might expect to observe a lagged decrease in N availability in response to increasing aridity. Nevertheless, our results suggest that the increase in aridity predicted with ongoing climate change will reduce N availability in the Mediterranean basin, impacting plant nutrient uptake and net primary production in semiarid grasslands throughout this region.

Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth’s land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.

Un estudio demuestra que los abetares más antiguos son más tolerantes a condiciones de aumento de temperatura y disminución de precipitación. El mantenimiento de la diversidad ayuda a los expertos a contar con individuos adaptados, idóneos en las restauraciones, que eviten el decaimiento de la especie.

Un estudio demuestra que los abetares más antiguos son más tolerantes a condiciones de aumento de temperatura y disminución de precipitación. El mantenimiento de la diversidad ayuda a los expertos a contar con individuos adaptados, idóneos en las restauraciones, que eviten el decaimiento de la especie.

The combined effects of shade and drought on plant performance and the implications for species interactions are highly debated in plant ecology. Empirical evidence for positive and negative effects of shade on the performance of plants under dry conditions supports two contrasting theoretical models about the role of shade under dry conditions: the trade-off and the facilitation hypotheses. We performed a meta-analysis of field and greenhouse studies evaluating the effects of drought at two or more irradiance levels on nine response variables describing plant physiological condition, growth, and survival. We explored differences in plant response across plant functional types, ecosystem types and methodological approaches. The data were best fit using quadratic models indicating a humped-back shape response to drought along an irradiance gradient for survival, whole plant biomass, maximum photosynthetic capacity, stomatal conductance and maximal photochemical efficiency. Drought effects were ameliorated at intermediate irradiance, becoming more severe at higher or lower light levels. This general pattern was maintained when controlling for potential variations in the strength of the drought treatment among light levels. Our quantitative meta-analysis indicates that dense shade ameliorates drought especially among drought-intolerant and shade-tolerant species. Wet tropical species showed larger negative effects of drought with increasing irradiance than semiarid and cold temperate species. Non-linear responses to irradiance were stronger under field conditions than under controlled greenhouse conditions. Non-linear responses to drought along the irradiance gradient reconciliate opposing views in plant ecology, indicating that facilitation is more likely within certain range of environmental conditions, fading under deep shade, especially for drought-tolerant species.

Las hojas son los órganos que usan las plantas en su interacción con la atmósfera, siendo determinantes en el balance de carbono y agua. Las características foliares tienen importantes consecuencias a nivel del funcionamiento de las plantas y de los ecosistemas, como por ejemplo en la producción primaria y en la circulación de nutrientes. Este trabajo se realizó en tres áreas protegidas del sur de la Península Ibérica (P. Natural Sierra de Cardeña y Montoro, P. Nacional de Sierra Nevada y P. Natural Los Alcornocales) en las que se midieron una serie de rasgos foliares en las especies leñosas más abundantes dentro de cada comunidad (28 especies en Cardeña, 19 en Sierra Nevada y 17 en Alcornocales). Los objetivos específicos fueron: 1) conocer como es la variación de estos rasgos foliares dentro de cada comunidad, 2) comprobar si alguno de los rasgos foliares estaban relacionados con diferentes estrategias, como la longevidad foliar (caducas vs. perennes) y formas de vida (árboles, matorrales y lianas) y 3) determinar si existen relaciones entre estos rasgos foliares y si las relaciones difieren entre los distintos sitios. Encontramos, en general, una gran variación en los rasgos foliares dentro de cada comunidad, como por ejemplo que el área foliar tuvo un coeficiente de variación del 100%. Se encontraron diferencias según la estrategia funcional, por ejemplo, las hojas caducas tenían mayor SLA y eran más finas que las perennes; y el contenido de materia seca de las hojas era menor en los matorrales y lianas con respecto a los árboles. Podemos concluir que, en el sur de la Península Ibérica, existe un espectro continuo de variación foliar originado por su diversidad filogenética y parcialmente determinado por la variación climática y las condiciones ambientales locales. Esta diversidad de estrategias puede contribuir a la coexistencia de especies dentro de la comunidad.

The objective of this study is to understand the response of Quercus suber seedlings to light and water, two limiting factors of their growth and survival in Mediterranean woody plants. Greenhouse and field experiments were conducted to contrast the results from both types of experiments. At the greenhouse experiment, Quercus suber seedlings were grown at three treatments of light (3%, 27% and 100% of incident light) and daily irrigation. When plants had four months old, half of the individuals did not receive any water, so we had then two water treatments (water and drought) within each light treatment. After two months, different variables were measured. At the field experiment, Quercus suber acorns were sown in three microhabitats with different light conditions (open, under tree and deep shade) in “Los Alcornocales” Natural Park (Cádiz). These microhabitats have similar light radiation than the treatments used at the greenhouse experiment. The greenhouse experiment results showed that the acorn mass was important for determining the seedling mass on the first stages, and that importance increased with light limitation. Thus, a greater seedling biomass was strongly associated with a bigger acorn mass under deep shade, but with increasing light levels the association between seedling and acorn mass became lower and non-significant. Quercus suber seedlings responded to light restriction with a decrease in the relative growth rate (RGR) and then the seedlings had lower biomass. This decrease in RGR was mainly caused by a decrease of the net assimilation rate (NAR). However, specific leaf area (SLA) and stem mass ratio (SMR) increased under shade as a way to enhance light capture. The response to water limitation was a general decrease of RGR, but interestingly, the effect of drought was negligible under deep shade conditions. The results of the field experiment were similar to those in the greenhouse experiment, which reinforce the conclusions of this study. Seedlin

Most of our knowledge regarding ontogenetic niche shifts in plants has been derived from studies involving only two or unconnected life stages. Approaches covering a broader range of different life stages are still needed to fully understand the implications of ontogenetic niche shifts for plant regeneration dynamics. We investigated ontogenetic shifts in the endangered Mediterranean tree species Acer opalus ssp. granatense (A. opalus) comparing the environmental characteristics of individuals of different ages with those of a random sample of available microsites. In addition, since herbivory could be a limiting factor, herbivory damage was quantified. Differences in environmental characteristics between locations of individual plants and randomly selected points became larger with plant age, suggesting that the regeneration niche of A. opalus shifts during ontogeny, undergoing a contraction. The presence of shrubs and adult trees, the depth of the litter layer, and herbivory were the main factors influencing these changes. A. opalus can germinate in all available microhabitats its seeds can reach, but saplings establish and grow only in a subset of microhabitats, which represent a change in tolerance to extrinsic factors. Old saplings establish under the canopy of shrubs, far away from tree cover that could block light required in the oldest stage. Therefore, temporal changes in the nature and strength of plant–plant interactions are also occurring. The ecological concept of niche shifts reveals the microsites with higher probabilities of long-term sapling survival of A. opalus: shrub cover involves an array of environmental changes crucial for the successful establishment of A. opalus saplings under stressful Mediterranean conditions, from microhabitat amelioration to herbivory protection.

[EN]: Growth rates of woody plant species: functional aspects and ecological implications. Plant species widely differ in their growth potential when they are grown under similar conditions. What are the factors related to a high growth potential? In general, a simple variable as the specific leaf area (the ratio between leaf area and leaf dry weight) is strongly and positively related to the relative growth rate (RGR) (explaining up to a 80% of variation in RGR). Also another factors as physiology (the net assimilation rate, NAR) can explain, although to a lesser extent, the differences in RGR between species. The biomass allocation to leaves is generally found to be positively related with RGR, whereas the allocation to roots is negatively related to RGR. The role of the architecture of the plant canopy is of increasing importance in plants of increasing size, but it has not been extensively studied. The biomass of a plant does not only depend on its RGR, but also on the initial biomass, which in the early stages is determined by the seed mass. In four Quercus species, the biomass of the seedlings was positively correlated with the seed mass. However, the importance of the seed mass was higher in shade conditions (about 3 % of incident light, a typical value in closed forest). The relative growth rates were also associated with the concentration of leaf chemical compounds. Protein concentration was positively correlated with RGR, but phenols and lignin concentration were negatively correlated with RGR. These results support the hypothesis of a trade-off between growth and defence: the energy is allocated to the synthesis of defence compounds, but at the expenses of a lower growth rate. In general, the characteristics associated with a higher growth capacity are those negatively related to seedling survivorship., [ES]: Tasas de crecimiento en especies leñosas: aspectos funcionales e implicaciones ecológicas. Las distintas especies vegetales difieren notablemente en su capacidad de crecimiento cuando se cultivan en condiciones parecidas. ¿Cuáles son las causas que determinan un mayor crecimiento? En general, una variable muy simple, a nivel foliar, como es el área específica foliar (la relación entre área y peso foliar) está estrecha y positivamente asociada con las tasas de crecimiento relativo (RGR) (explicando hasta un 80% de la variación de RGR). También otros factores como la fisiología (la tasa de asimilación neta, NAR) puede explicar, aunque en menor grado, las diferencias en RGR entre especies. La distribución de biomasa a las hojas también se encuentra generalmente relacionada positivamente con RGR, mientras que la distribución a raíces lo está negativamente. El papel de la arquitectura de la copa parece que puede tener una importancia mayor en plantas de mayor tamaño. Por otro lado, el peso de una planta en un momento determinado, no solo depende de su RGR sino también del peso inicial, que viene determinado por el peso de la semilla. En cuatro especies de Quercus, el peso de la plántula estaba correlacionado con el peso de la semilla. Sin embargo, la importancia de la semilla era mucho mayor en condiciones de luz limitante (alrededor de un 3%, típica de ambientes de bosque cerrado). Las tasas de crecimiento relativo también estaban asociadas a la concentración de compuestos químicos de la hoja. La concentración de proteínas estaba correlacionada positivamente con RGR, mientras que la concentración de fenoles solubles y lignina lo estaban negativamente. Estos resultados apoyarían la hipótesis de compromiso entre crecimiento y defensa, de forma que la energía que la planta dedica a sintetizar compuestos defensivos es a costa de un menor crecimiento y viceversa. De forma general, las características asociadas a una mayor capacidad de crecimiento son aquellas que están

Understanding the impacts of combined resource supplies on seedlings is critical to enable prediction of establishment growth, and forest dynamics. We investigated the effects of irradiance and water treatments on absolute growth, and relative growth rate (RGR) and its components, for seedlings of four Quercus species differing in leaf habit and with a wide variation in seed mass. Plants were grown for 6.5 months at three levels of irradiance (100, 27, and 3% daylight), and treated during the last 2.5 months with two watering treatments (frequent watering v. suspended watering). Both shade and drought reduced seedling growth rates, with a significant interaction: under full irradiance the drought treatment had a stronger impact on RGR and final biomass than under deep shade. For three species, seed mass was positively related to absolute growth, with stronger correlations at lower irradiance. The evergreen species grew faster than the deciduous species, though leaf habit accounted for a minor part of the interspecific variation in absolute growth. Seedling biomass was determined positively either by RGR or seed mass; RGR was positively linked with net assimilation rate (NAR) and leaf mass fraction (LMF), and seed mass was negatively linked with RGR and LMF, but positively linked with NAR. Seedling RGR was not correlated with light-saturated net photosynthetic rate, but was strongly correlated with the net carbon balance estimated, from photosynthetic light-response curves, considering daily variation in irradiance. These findings suggest an approach to applying short-term physiological measurements to predict the RGR and absolute growth rate of seedlings in a wide range of combinations of irradiance and water supplies.

[EN]: Light and water availability are two of the main factors that regulate growth and survival of Mediterranean woody plants. The objective of this study is to investigate the plastic response of seedlings of four Mediterranean Quercus species to these factors, as well as to determine what traits are more plastic to one or another factor. Our hypothesis was that physiological variables are more plastic than structural variables. We cultivated in controlled conditions seedlings of Quercus ilex ssp. ballota, Q. suber, Q. pyrenaica and Q. canariensis in three treatments of light (100, 27 and 3 % of incident light) and with abundant water. On late spring, watering was stopped for half of the seedlings, so there were two water treatments (irrigation versus no-irrigation) within each treatment of light. Several physiological traits were studied: photosynthetic and respiration rate, light compensation and saturation points, stomatal conductance, water use and nitrogen use efficiency. We also evaluated different structural traits were studied such as leaf area, specific leaf area, leaf nitrogen and carbon concentrations, and a chlorophyll index. Changes in water availability induced low plastic responses of structural traits, whereas responses of physiological traits were relatively higher. On the other hand, there were not significant differences in the plastic response of structural or physiological traits to changes in light availability. In these Mediterranean oak species, the higher response to light could be linked to a shade tolerance strategy and the low response to water (i.e., canalization) could be related to drought tolerant strategy, thus indicating that tolerance to low-resources may be attained through different strategies., [ES]: La luz y el agua son dos de los principales factores que regulan el crecimiento y la supervivencia de las plantas leñosas mediterráneas. El objetivo de este estudio es conocer el nivel de respuesta plástica de plántulas de cuatro especies de Quercus a tratamientos de sombra y sequía. Además queremos evaluar qué variables son más plásticas a uno u otro factor, y probar la hipótesis de que los rasgos fisiológicos son más plásticos que los rasgos estructurales. Se cultivaron en condiciones controladas plántulas de Quercus ilex subsp. ballota, Q. suber, Q. pyrenaica y Q. canariensis en tres tratamientos de luz (100, 27 y 3 % de luz incidente) y con agua abundante. A partir de mediados de primavera, a la mitad de las plántulas se les suprimió el agua, de forma que recibieron dos tratamientos de agua (riego versus no riego) dentro de cada tratamiento de luz. Se analizaron distintos rasgos fisiológicos como la tasa de fotosíntesis y respiración, el punto de compensación y de saturación a la luz, la conductancia estomática, la eficiencia en el uso del agua y de nitrógeno y la eficiencia fotosintética a la luz. También se estudiaron distintos rasgos morfológicos o estructurales, como el área foliar, el área específica foliar, las concentraciones foliares de N y C, y un índice de clorofila. En general, la respuesta plástica de las variables estructurales al factor agua fue muy baja, mientras que la respuesta de las fisiológicas fue relativamente alta. Con la luz, no se detectaron diferencias significativas en la respuesta plástica entre las variables estructurales y fisiológicas. En estas especies de Quercus mediterráneos, la mayor respuesta plástica a la luz podría estar ligada con una estrategia de tolerancia a la sombra, mientras que la baja respuesta plástica al agua (i.e., canalización) respondería a una estrategia de tolerancia a la sequía, indicando que la tolerancia a bajos niveles de recursos puede ser alcanzada mediante diferentes estrategias.

Understanding seedling performance across resource gradients is crucial for defining the regeneration niche of plant species under current environmental conditions and for predicting potential changes under a global change scenario. A 2-year field experiment was conducted to determine how seedling survival and growth of two evergreen and two deciduous Quercus species vary along gradients of light and soil properties in two Mediterranean forests with contrasting soils and climatic conditions. Half the seedlings were subjected to an irrigation treatment during the first year to quantify the effects on performance of an alteration in the summer drought intensity. Linear and non-linear models were parameterized and compared to identify major resources controlling seedling performance. We found both site-specific and general patterns of regeneration. Strong site-specificity was found in the identity of the best predictors of seedling survival: survival decreased linearly with increasing light (i.e. increasing desiccation risk) in the drier site, whereas it decreased logistically with increasing spring soil water content (i.e. increasing waterlogging risk) in the wetter site. We found strong empirical support for multiple resource limitation at the drier site, the response to light being modulated by the availability of soil resources (water and P). Evidence for regeneration niche partitioning among Quercus species was only found at the wetter site. However, at both sites Quercus species shared the same response to summer drought alleviation through water addition: increased first-year survival but not final survival (i.e. after two years). This suggests that extremely dry summers (i.e. the second summer in the experiment) can cancel out the positive effects of previous wetter summers. Therefore, an increase in the intensity and frequency of summer drought with climate change might cause a double negative impact on Quercus regeneration, due to a general reduction in survi

Regeneration ecology of mixed Quercus suber (cork oak) - Q. canariensis (Algerian oak) forests of southern Spain has been studied at different scales and levels of ecological organization. We present results on: 1) species regeneration patterns at regional scale, based on the Spanish Second Forest Inventory data; 2) environmental heterogeneity found at forest stand scale; 3) acorn removal rates along a gradient of canopy cover (light availability) as function of seed size, based on field experiments; 4) patterns of seed germination, and seedling emergence and survival along a combined gradient of light and water availability, based on seed sowing experiments; 5) differential response of oak species to combined treatments of shade and drought, under greenhouse conditions; 6) models predicting oaks’ regeneration patterns under different environmental conditions.

Three hypotheses have been proposed to explain the functional relationship between seed mass and seedling performance: the reserve effect (larger seeds retain a larger proportion of reserves after germinating), the metabolic effect (seedlings from larger seeds have slower relative growth rates), and the seedling-size effect (larger seeds produce larger seedlings). We tested these hypotheses by growing four Mediterranean Quercus species under different light conditions (3, 27, and 100% of available radiation). We found evidence for two of the three hypotheses, but none of the four species complied with all three hypotheses at the same time. The reserve effect was not found in any species, the metabolic effect was found in three species (Q. ilex, Q. pyrenaica, and Q. suber), and the seedling-size effect in all species. Light availability significantly affected the relationships between seed size and seedling traits. For Q. ilex and Q. canariensis, a seedling-size effect was found under all three light conditions, but only under the lowest light (3%) for Q. suber and Q. pyrenaica. In all species, the correlation between seed mass and seedling mass increased with a decrease in light, suggesting that seedlings growing in low light depend more upon their seed reserves. A causal model integrates the three hypotheses, suggesting that larger seeds generally produced larger seedlings.

SADIE es una herramienta de análisis espacial basado en índices de distancia. Su sencilla metodología, cuantifica el patrón espacial de una variable objeto de estudio, detecta los agregados locales de la variable y cuantifica el grado de asociación o disociación espacial entre pares de variables. SADIE genera estadísticos continuos para cada punto de muestro con lo que las diferentes facetas del patrón espacial que SADIE detecta, puede ser representadas mediante mapas. Otra de las ventajas de SADIE es su baja exigencia con datos de diferente naturaleza aunque también se describen diferentes limitaciones de la metodología. El creador de SADIE también ha diseñado un sencillo programa informático, bajo entorno Windows, que facilita la utilización de esta herramienta. A modo de ejemplo, se presentan resultados de un estudio sobre la variación espacial a pequeña escala de factores abióticos en la heterogénea Sierra Nevada (Granada). La versatilidad y sencillez de uso, han hecho que esta herramienta de análisis espacial se haya extendido considerablemente en los últimos tiempos., SADIE is a tool for spatial analysis based in distance indexes. Its simply methodology, quantifies the spatial pattern of a focal variable, reveals local clusters of the variable and measures the association or dissociation degree between variables. SADIE creates continuous statistics in each sample point, in this way, different characteristics of the spatial pattern should be mapped. SADIE can be use with data of different nature although different shortcomings of this methodology are described. The creator of SADIE have also designed a software, which can be use in windows, facilitating the harness of this tool. As example, different results from a field study about small-scale spatial variation of a biotic factors in Sierra Nevada National Park are presented. Lately, this spatial analysis tool has been widely used in scientific literature because its versatility and simplicity.

SADIE es una herramienta de análisis espacial basado en índices de distancia. Su sencilla metodología, cuantifica el patrón espacial de una variable objeto de estudio, detecta los agregados locales de la variable y cuantifica el grado de asociación o disociación espacial entre pares de variables. SADIE genera estadísticos continuos para cada punto de muestro con lo que las diferentes facetas del patrón espacial que SADIE detecta, puede ser representadas mediante mapas. Otra de las ventajas de SADIE es su baja exigencia con datos de diferente naturaleza aunque también se describen diferentes limitaciones de la metodología. El creador de SADIE también ha diseñado un sencillo programa informático, bajo entorno Windows, que facilita la utilización de esta herramienta. A modo de ejemplo, se presentan resultados de un estudio sobre la variación espacial a pequeña escala de factores abióticos en la heterogénea Sierra Nevada (Granada). La versatilidad y sencillez de uso, han hecho que esta herramienta de análisis espacial se haya extendido considerablemente en los últimos tiempos., SADIE is a tool for spatial analysis based in distance indexes. Its simply methodology, quantifies the spatial pattern of a focal variable, reveals local clusters of the variable and measures the association or dissociation degree between variables. SADIE creates continuous statistics in each sample point, in this way, different characteristics of the spatial pattern should be mapped. SADIE can be use with data of different nature although different shortcomings of this methodology are described. The creator of SADIE have also designed a software, which can be use in windows, facilitating the harness of this tool. As example, different results from a field study about small-scale spatial variation of a biotic factors in Sierra Nevada National Park are presented. Lately, this spatial analysis tool has been widely used in scientific literature because its versatility and simplicity.

Here, we investigated the physiological and structural leaf responses of seedlings of two evergreen and two deciduous Quercus species, grown in a glasshouse and subjected to contrasted conditions of light (low, medium and high irradiance) and water (continuous watering vs 2-months drought). • The impact of drought on photosynthetic rate was strongest in high irradiance, while the impact of shade on photosynthetic rate was strongest with high water supply, contradicting the hypothesis of allocation trade-off. • Multivariate causal models were evaluated using d-sep method. The model that best fitted the dataset proposed that the variation in specific leaf area affects photosynthetic rate and leaf nitrogen concentration, and this trait determines stomatal conductance, which also affects photosynthetic rate. • Shade conditions seemed to ameliorate, or at least not aggravate, the drought impact on oak seedlings, therefore, the drought response on leaf performance depended on the light environment.

Field experiments reporting the relative growth rate (RGR) patterns in plants are scarce. In this study, 22 herbaceous species (20 Aegilops species, Amblyopyrum muticum and Triticum aestivum) were grown under field conditions to assess their RGR, and to find out if the differences in RGR amongst species were explained by morphological or physiological traits. Plants were cultivated during two months, and five harvests (every 13–19 days) were carried out. Factors explaining between-species differences in RGR varied, depending on whether short (13–19 days) or longer periods (62 days) were considered. RGR for short periods (4 growth periods of 13–19 days each) showed a positive correlation with net assimilation rate (NAR), but there was no significant correlation with leaf area ratio (LAR) (with the exception of the first growth period). In contrast, when growth was investigated over two months, RGR was positively correlated with morphological traits (LAR, and specific leaf area, SLA), but not with physiological traits (NAR). A possible explanation for these contrasting results is that during short growth periods, NAR exhibited strong variations possibly caused by the variable field conditions, and, consequently NAR mainly determined RGR. In contrast, during a longer growth period (62 days) the importance of NAR was not apparent (there was no significant correlation between RGR and NAR), while allocation traits, such as LAR and SLA, became most relevant.

Se presentan resultados preliminares del crecimiento de plántulas en un experimento de campo donde se sembraron semillas en diferentes tipos de micrositios con distinta intensidad luminosa (abierto, bajo árbol y sombra densa), y en un experimento de invernadero donde se aplicaron trata- mientos de luz y riego. En el campo, las plántulas de Q. suber de los micrositios abiertos fueron mayo- res que bajo los árboles. En contraste, la supervivencia fue menor en los mismos sitios abiertos, en comparación con la zona bajo los árboles. La zona de sombra densa fue la más adversa: además de ser las plántulas más pequeñas, la supervivencia fue la menor. Existe, por tanto, un desacoplamiento entre la supervivencia y el crecimiento de las plántulas de Quercus. En el invernadero, los niveles más altos de luz y de agua determinaron una mayor biomasa de las plántulas, pero las especies respondie- ron de forma diferente. Las plántulas de Q. suber tuvieron mayor biomasa que las de Q. canariensis. Se encontró una interacción entre los tratamientos de luz y agua: en la sombra densa no se manifes- taron diferencias entre el peso de las plántulas regadas respecto a las que sufrieron la sequía.

Se presentan algunos resultados sobre la ecología de la regeneración de las comunidades de bosques dominados por especies de Quercus (especialmente Q. suber y Q. canariensis) en el sur de España. "La ecología de la regeneración es un proceso múltiple, que incluye la floración y fructificación de los árboles maduros y la dispersión de sus semillas; así como la germinación y establecimiento de las plántulas, su crecimiento y los cambios poblacionales de los árboles (Quercus) y de las plantas asociadas" (Johnson, Shifley y Rogers, 2002). La comunidad vegetal del bosque es bastante diversa; parte de esta diversidad está almacenada en los bancos de semillas (p.ej., 11.260 semillas/m2 de 28 especies en el alcornocal).

Spatial heterogeneity of abiotic factors influences patterns of seedling establishment at different scales. In stress-prone ecosystems such as Mediterranean ones, heterogeneity generated by shrubs has been shown to facilitate the establishment of tree species. However, how this facilitation is affected by spatial scale remains poorly understood. We have experimentally analysed the consequences of the abiotic heterogeneity generated by pioneer shrubs on survival, growth and physiology of seedlings of three important tree species from Mediterranean mountains (Acer opalus ssp. granatense, Quercus pyrenaica and Quercus ilex). Patterns of abiotic heterogeneity and seedling performance were studied at two scales differing in grain: 1) the microhabitat scale, by using open interspaces as controls of the effect of different shrub species, and 2) the microsite scale, analysing the effects of fine-grain heterogeneity (within-microhabitat heterogeneity). Results showed that, at the microhabitat scale, seedling establishment of the three tree species significantly benefited from the modification of the abiotic environment by nurse shrubs. However, we found shrub-seedling interactions to be species-specific, due to differential modification of both aboveground (light availability) and belowground (soil compaction, water content, and fertility) abiotic factors by nurse shrub species. Heterogeneity at the within-microhabitat scale was rather high, although it did not significantly affect seedling performance of any of the tree species. The study demonstrates that the effects of the abiotic heterogeneity generated by shrubs are not consistent across the range of spatial scales considered. The regeneration niche of tree species becomes very complex at fine spatial scales, and thereby estimators of abiotic heterogeneity are valuable descriptors of spatial patterns of seedling establishment only when microsite ‘‘noise’’ is averaged out at greater scales.

Understanding the processes of tree population recruitment and their limitations, is the scientific basis to assure the forest natural regeneration, and to improve techniques of restoration and afforestation. We present here preliminary results of a collaborative project (HETEROMED). The environmental heterogeneity of Mediterranean forests was related to natural patterns of seedling establishment. Factorial design experiments of seed addition were carried out and the resulting emergence and survival were related to the conditions of light and soil moisture. The fieldwork was carried out at two natural areas in South Spain: Sierra Nevada National Park (higher mountains on the Southeast) and Los Alcornocales Natural Park (lower mountains in Sierra del Aljibe range, on the southern tip of the Iberian Peninsula). The experiments were focussed in seven target species - Quercus suber, Quercus canariensis, Quercus ilex, Quercus pyrenaica, Pinus sylvestris, Acer granatense and Sorbus aria. In addition, a glasshouse factorial experiment investigated the differential response of the four Quercus species to the combined effects of shade and drought. In both field sites (Sierra Nevada and Sierra del Aljibe) seedling emergence differed among species, but was not significantly affected by habitat (light availability). In Sierra Nevada, seedling survival after dry summer was statistically different between tree species, habitat type and irrigation treatment. Summer drought was the main cause of seedling mortality. Survival was higher in deep shade habitats than in open stands. Irrigation boosted survival in open areas (full light) and under shrublands (medium shade), but had scant effect inside woodland stands (deep shade). In Sierra del Aljibe, there were significant differences in seedling survival of Quercus suber between habitat types. Most surviving seedlings after the dry summer were in habitats with medium shade (under tree canopy), rather than in full light (open areas) or