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* Background and Aims In Mediterranean annual plants, germination mainly occurs during the autumn and only those seedlings that survive winter freezing can flower and produce seedlings in spring. Surprisingly, the effect of freezing events as an abiotic determinant of these communities remains unexplored. The present study aimed to investigate how freezing events affect annual Mediterranean communities and whether their functional structure as related to freezing resistance is linked to the main biotic and abiotic determinants of these communities. * Methods In 120 plots located on a semi-arid Mediterranean steppe (Spain), the community functional structure related to the lethal temperature causing 50 % frost damage (LT50 trait) in seedlings was estimated and summarized as the community-weighted mean (CWM-LT50) and its functional diversity (FD-LT50). Plots were stratified according to distance to rabbit shelters and latrines as a proxy for rabbit density, proximity to Stipa tenacissima and spring water availability, where annual species abundance was recorded in all plots over three consecutive years. * Key Results Annual species were able to resist a threshold temperature of -4 °C and most had LT50 values around the absolute minimum temperature (-9.5 °C) in the three years. Higher rabbit densities led to lower CWM-LT50 and higher FD-LT50 values. Plots close to Stipa tussocks had higher CWM-LT50 values whereas water availability had no effects. * Conclusions High freezing resistance was extended among winter annual species, suggesting the presence of an association between historical environmental filtering and low winter temperatures. However, the community functional structure related to freezing resistance remained variable among scenarios with differences in herbivory pressure and distance to perennial vegetation. The trends observed indicate that traits that allow plants to deal with herbivory may also promote freezing resistance, and that tussocks can act as nurses via microclimatic amelioration of harsher winter conditions. [ABSTRACT FROM AUTHOR]

Verticillium wilt in olive is considered the most serious biotic threat to this crop. Effective control of this disease relies on an integrated disease management strategy in which the biological control has an important role nowadays. This work describes the potential effect of 44 plant extracts and 20 essential oils against Verticillium dahliae. The results demonstrate the in vitro and in planta effectiveness of essential oil from Thymus, in particular Thymus sp. 04 (prepared in the laboratory), and the commercial product Thymus sp. 01, against Verticillium dahliae. The inhibition of mycelial growth and microsclerotia reached 100% in both treatments and achieved a disease reduction in olive plants by 65% and 42% for Thymus sp. 04 and sp. 01, respectively. These treatments showed the potential for essential oils use in the control of this pathogen in the frame of an integrated disease management strategy. This is the first report of the use of essential oils to control Verticillium wilt in olive plants. Further studies are warranted to identify the bioactive compounds in the essential oil that control V. dahliae and evaluate their potential use as natural fungicides. [ABSTRACT FROM AUTHOR]

Background and Aims Many studies have analysed the mechanisms that determine plant coexistence in standing vegetation, but the determinants of soil seed bank species assemblies have rarely been studied. In gypsum soil communities, aerial vegetation and seed banks are tightly connected in space and time, but the mechanisms involved in their organization may differ. The aim of this study is to understand the relative importance of biotic and abiotic factors controlling soil seed bank composition and structure. Methods Persistent and complete (i.e. persistent plus transient) soil seed banks were investigated at two spatial scales in a very species-rich semi-arid community dominated by annuals. A water addition treatment equivalent to 50% annual increase in average precipitation (abiotic factor) was applied for two consecutive years, and the relationships of the soil seed bank to the biological soil crust (BSC), above-ground vegetation and the presence of Stipa tenacissima tussocks (biotic factors) were simultaneously evaluated. Key Results As expected, the standing vegetation was tightly related to seed abundance, species richness and composition in both seed banks. Remarkably, BSC cover was linked to a decrease in seed abundance and species richness in the persistent seed bank, and it even determined complete seed bank composition at the fine spatial scale. However, this effect disappeared at coarser scales, probably because of the high spatial heterogeneity induced by BSCs. In contrast to findings on standing vegetation, Stipa and the irrigation treatment for two consecutive years had no effect on soil seed banks. Conclusions Soil seed bank assemblies in our semi-arid plant community were the result of above-ground vegetation dynamics and of the direct filtering processes on seed fate operated by the spatially heterogeneous BSCs. Cover of BSCs was negatively correlated with seed abundance and species richness, and affected seed species composition in the soil. Changes in species composition and enrichment when the BSC cover is low suggest that BSCs promote a fine scale niche differentiation in the soil seed bank and thereby potentially enhance species coexistence and high species diversity in these communities. [ABSTRACT FROM AUTHOR]

Assessing freezing community response and whether freezing resistance is related to other functional traits is essential for understanding alpine community assemblages, particularly in Mediterranean environments where plants are exposed to freezing temperatures and summer droughts. Thus, we characterized the leaf freezing resistance of 42 plant species in 38 plots at Sierra de Guadarrama (Spain) by measuring their ice nucleation temperature, freezing point (FP), and low-temperature damage (LT50), as well as determining their freezing resistance mechanisms (i.e., tolerance or avoidance). The community response to freezing was estimated for each plot as community weighted means (CWMs) and functional diversity (FD), and we assessed their relative importance with altitude. We established the relationships between freezing resistance, growth forms, and four key plant functional traits (i.e., plant height, specific leaf area, leaf dry matter content (LDMC), and seed mass). There was a wide range of freezing resistance responses and more than in other alpine habitats. At the community level, the CWMs of FP and LT50 responded negatively to altitude, whereas the FD of both traits increased with altitude. The proportion of freezing-tolerant species also increased with altitude. The ranges of FP and LT50 varied among growth forms, and only leaf dry matter content was negatively correlated with freezing-resistance traits. Summer freezing events represent important abiotic filters for assemblies of Mediterranean high mountain communities, as suggested by the CWMs. However, a concomitant summer drought constraint may also explain the high freezing resistance of species that thrive in these areas and the lower FD of freezing resistance traits at lower altitudes. Leaves with high dry matter contents may maintain turgor at lower water potential and enhance drought tolerance in parallel to freezing resistance. This adaptation to drought seems to be a general prerequisite for plants found in xeric mountains. [ABSTRACT FROM AUTHOR]

Climate change is expected to alter the geographic distribution of many plant species worldwide. However, there is still no clear evidence showing a generalised direction and magnitude of these distribution shifts. Here, we have surveyed, in nine mountainous regions in Spain, an array of tree species along entire elevational ranges, as surrogates of their global climatic ranges, to test for elevational shifts towards cooler locations. We analysed the distribution recruitment patterns of five dominant tree species, recording the abundance and measuring the primary growth of juveniles in 306 plots. Three of the species have a temperate-boreal distribution with populations at their southern edge in the Mediterranean mountain ranges: Pinus sylvestris , Pinus uncinata and Fagus sylvatica ; and the other two species have a Mediterranean distribution: Quercus ilex and Pinus nigra . Despite the contrasting phylogenies and biogeographies, we identified a similar pattern in recruitment abundance across species, with an asymmetric distribution of juveniles (more recruits in the middle-upper elevation of their range), but higher annual growths at lower elevations. This survival-growth trade-off at the early recruitment stage may potentially counterbalance at population level the negative effect of global warming on recruit survival at the lower edge of species ranges. These findings suggest a demographic stabilisation process at the early recruitment stage of these tree species, and highlight the importance of considering the different demographic stages across the whole climatic range to understand the effects that climate change may exert on species distributions and population dynamics. [ABSTRACT FROM AUTHOR]

Cold stratification provided by snow cover is essential to break seed dormancy in many alpine plant species. The forecast reduction in snow precipitation and snow cover duration in most temperate mountains as a result of global warming could threaten alpine plant populations, especially those at the edge of their species distribution, by altering the dynamics of early life stages. We simulated some effects of a reduction in the snow cover period by manipulating the duration of cold stratification in seeds of Silene ciliata, a Mediterranean alpine specialist. Seeds from three populations distributed along an altitudinal gradient were exposed to different periods of cold stratification (2, 4 and 6 months) in the laboratory and then moved to common garden conditions in a greenhouse. The duration of the cold stratification treatment and population origin significantly affected seed emergence percentage, emergence rate and seedling size, but not the number of seedling leaves. The 6-month and 4-month cold stratification treatments produced higher emergence percentages and faster emergence rates than seeds without cold stratification treatment. No significant cold stratification duration x seed population origin interactions were found, thus differential sensitivity to cold stratification along elevation is not supported. [ABSTRACT FROM AUTHOR]

Assessing changes in plant functional traits along gradients is useful for understanding the assembly of communities and their response to global and local environmental drivers. However, these changes may reflect the effects of species composition (i.e. composition turnover), species abundance (i.e. species interaction), and intra-specific trait variability (i.e. species plasticity). In order to determine the relevance of the latter, trait variation can be assessed under minimal effects of composition turnover. Nine sampling sites were established along an altitudinal gradient in a Mediterranean high mountain grassland community with low composition turnover (Madrid, Spain; 1940 m–2419 m). Nine functional traits were also measured for ten individuals of around ten plant species at each site, for a total of eleven species across all sites. The relative importance of different sources of variability (within/between site and intra-/inter-specific functional diversity) and trait variation at species and community level along the considered gradients were explored. We found a weak individual species response to altitude and other environmental variables although in some cases, individuals were smaller and leaves were thicker at higher elevations. This lack of species response was most likely due to greater within- than between-site species variation. At the community level, inter-specific functional diversity was generally greater than the intra-specific component except for traits linked to leaf element content (leaf carbon content, leaf nitrogen content, δ13C and δ15N). Inter-specific functional diversity decreased with lower altitude for four leaf traits (specific leaf area, leaf dry matter content, δ13C and δ15N), suggesting trait convergence between species at lower elevations, where water shortage may have a stronger environmental filtering effect than colder temperatures at higher altitudes. Our results suggest that, within a vegetation type encompassing various environmental gradients, both, changes in species abundance and intra-specific trait variability adjust for the community functional response to environmental changes. [ABSTRACT FROM AUTHOR]

In harsh environments facilitation alleviates biotic and abiotic constraints on tree recruitment. Under ongoing drier climate change, we expect facilitation to increase as a driver of coexistence. However, this might not hold under extreme abiotic stress and when the outcome depends on the interaction with other drivers such as altered herbivore pressure due to land use change. We performed a field water-manipulation experiment to quantify the importance of facilitation in two coexisting Mediterranean trees (dominant Juniperus thurifera and coexisting Quercus ilex subsp. ballota) under a climate change scenario. Shifts in canopy dominance favouring Q. ilex could be based on the extension of heterospecific facilitation to the detriment of conspecific alleviation. We found that saplings of both species transplanted under the canopy of nurse trees had greater survival probability, growth and photochemical efficiency. Intra- and interspecific facilitation mechanisms differed: alleviation of abiotic stress benefited both species during summer and J. thurifera during winter, whereas browsing protection was relevant only for Q. ilex. Facilitation was greater under the dry treatment only for Q. ilex, which partially agreed with the predictions of the stress gradient hypothesis. We conclude that present rainfall availability limits neither J. thurifera nor Q. ilex establishment. Nevertheless, under current global change scenarios, imposing increasing abiotic stress together with altered herbivore browsing, nurse trees could differentially facilitate the establishment of Q. ilex due to species-specific traits, i.e. palatability; drought, heat and cold tolerance, underlying species differences in the facilitation mechanisms and eventually triggering a change from pure juniper woodlands to mixed formations. [ABSTRACT FROM AUTHOR]

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species ( Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems. [ABSTRACT FROM AUTHOR]

Background and aims: Many studies have revealed marked changes in the patterns of nitrogen use along ontogeny in tree species. However, there are no reports of the ontogenetic changes in resorption efficiency, despite the fact that nitrogen resorption from senescing leaves is one of the most important plant mechanisms to conserve and optimize the use of this nutrient. Our objective is to analyze the differences in nitrogen resorption between seedlings and adults and to elucidate which of the leaf traits changing along ontogeny determine these possible differences. Methods: Green and senescent leaves were sampled from five seedlings and mature specimens of four tree species. N resorption efficiencies were calculated from the differences between N contents in green and shed leaves collected from the same specimens. Minimum N concentrations in leaf litter were used as an estimation of resorption proficiency. Results: N concentration in green leaves and N resorption efficiency were higher in adult specimens for all species. However, with the exception of Pinus pinea, there were no significant differences in proficiency. Conclusions: Contrary to our expectations, the species studied seem to have a lower potential for N resorption during the initial stages of their life cycle. The higher N concentrations seen in adult leaves could explain the higher percentage they are able to withdraw during senescence, probably because they have higher amounts of readily translocatable soluble molecules. [ABSTRACT FROM AUTHOR]

Questions What changes have occurred in Mediterranean high-mountain communities during the last 50 yr? Do these changes reflect upward shifts and decline in dry grassland specialists due to climate warming? Location Upper belt (1900-2400 m a.s.l.) in Sierra de Guadarrama, Spain. Methods We re-surveyed dry grassland communities in 50 summits that were previously sampled between the 1960s and 1980s. New plots were placed in the same localities and the same conditions as the original surveys. Changes at the species level were evaluated by comparing, between the two survey periods, species frequencies and optimal altitude estimated by local distribution models done with Maxent. Changes at the community level were assessed by comparing species richness and composition of the two surveys in relation to altitude, plot size and time between surveys. We evaluated observed changes in the whole data set and species groups by structuring the species into dry grassland specialists, other alpine species and generalists occurring at wider altitudinal ranges. Results There was a general decline in the frequency of dry grassland specialists, an increase in frequency of generalists and no clear trends in the other species. Upward shifts were mainly detected in generalists, with significant increases in frequency. At the community level, we found a general increase in species richness that was mainly determined by the increase in generalists; coupled with a decline in number of dry grassland specialists. The structure of the communities evidenced changes between the historical surveys and the re-survey, which were mainly correlated with the number of years and the increasing dominance of generalists. Conclusions In contrast with trends observed in other Mediterranean high-mountain communities, our study reports a general increase in species richness, although this effect is concurrent with a decline in dry grassland specialists. The observed trends support on-going replacement of endemic species by low-altitude species, and associated changes in community assemblages. Despite the limitations of re-visitation studies for assessing climate-driven changes, our results highlight the necessity of developing more studies for assessing the effects of drought in Mediterranean high-mountain communities. [ABSTRACT FROM AUTHOR]

Whether or not it is time for germination is a dilemma for annuals since they have only a single opportunity to reproduce successfully. This is critical for species living in stressful and unpredictable environments such as Mediterranean habitats. In order to clarify the environmental cues of germination, four annuals of different families, all of them occurring very frequently on gypsum soils, were selected and their germination observed under 14 climate scenarios, mimicking temperature and photoperiod conditions from autumn to spring, at two levels of water availability (continuous irrigation versus an initial single irrigation event followed by a progressive soil desiccation). In spring scenarios, two seed-storage conditions were compared: dry cold and room conditions. In the absence of water limitation, germination decreased from early autumn to late spring. Water scarcity always reduced germination, especially in early spring. Our results suggest a facultative winter germination behaviour and highlight the crucial role of dry cold storage in reducing spring germination. In conclusion, Mediterranean ephemerals showed a very plastic germination response that allows them to take advantage of favourable environmental conditions from autumn to spring. This environmental cueing is combined with the ability to dilute the risk through a variable rate of seed dormancy that, according to bet-hedging strategies, increases from secure autumn to riskier spring. [ABSTRACT FROM PUBLISHER]

Abstract: The holocentric structure promotes chromosome rearrangements by fission, fusion, translocation, and inversion, which have been thought to promote differentiation and speciation. The Carex laevigata group (Cyperaceae) comprises four species: two restricted endemics from the western Mediterranean (Carex camposii, 2n =72, and Carex paulo-vargasii, 2n =74–75), and two more widespread species, found mostly in Western Europe (C. laevigata, 2n =69–84, and Carex binervis, 2n =72–74). We tested the selection for chromosome number by climatic variables by controlling for the non-independence of the data using generalized linear mixed model (GLMM). We obtained chromosome counts as well as DNA sequences for the 5′trnK intron and the trnV-ndhC intergenic spacer in the chloroplast genome from 181 individuals from 53 populations representing these four species. We also climatically characterized the sites where the 53 populations were found using the WorldClim database. Our results show that the best predictor of chromosome number variation is the climatic environment rather than neutral evolutionary processes like founder events and migration patterns. These results support the adaptive value of the holocentric chromosomes and their role in promoting differentiation and eventually speciation. [Copyright &y& Elsevier]

Abstract: Evaluation of the effectiveness of protected areas networks for conserving diversity is a demanding topic in biological conservation. In the last few years, there has been an increasing number of studies on species distribution modelling which can be easily used to test the efficiency of reserve networks. Unfortunately, this effort has been focused on the so called charismatic species. Our main goal is to test the effectiveness of the Natura 2000 network by using non-charismatic species. We determined potential distribution patterns for 18 lichen species in Spain and evaluated the effectiveness of the Natura 2000 network in protecting them. Models were implemented with ENFA (Ecological-Niche Factor Analysis) because this modelling approach only requires presence data since the reliability of absence data is really poor. Habitat suitability maps were obtained and high suitability areas were characterised using ten environmental predictors. Maps of areas of high suitability were overlaid with the Natura 2000 network cover, considering the different biogeographical regions present in mainland Spain. The effectiveness of the Natura 2000 was quite low in most species. The analysis per region showed that Mediterranean lichen species growing in forests are better protected than species found in coastal, drier and warmer areas. Our results indicate that the Natura 2000 network, a protection system based on vascular plants, does not guarantee the protection of Mediterranean lichen species. Thus, this type of gap analysis and especially the inclusion of “non-charismatic” organisms such as lichens should be considered in studies to define protected areas. [Copyright &y& Elsevier]

• Premise of the study: Regional climatic patterns result in the synchrony of biological processes along large spatial areas. These patterns may be critical for effective plant recruitment in (semi)arid environments. Nevertheless, recruitment patterns of plant species within their range of distribution are still poorly known. Moreover, this response may be species-dependent, particularly between coexisting congenerics, which are thought to vary in demographic responses to climatic conditions as a coexistence-promoting mechanism. In this context, we investigated whether two congeneric plant species show synchronic age structures at varying spatial scales and whether they display demographic differences. • Methods: We sampled 600 Helianthemum squamatum and H. syriacum individuals at six localities in Spain. We used dendro-chronological techniques to estimate age and radial growth. We compared the age structure among populations and between species and assessed the effect of site and species on different demographic parameters. Correlations between age structure and climatic factors were also determined. • Key results: We detected a very low intraspecific synchrony across sites but a high interspecific coupling in age structure within localities. Secondary growth, mean age, and flowering showed large intersite differences and small interspecific differences. Summer rainfall was a good predictor of age structure. • Conclusions: Fine-scale climatic variability plays a major role in determining age structure of the studied species. Climatic variability is more relevant than interspecific differences. Our results illustrate the relevance of including spatial variation in summer rainfall patterns when modeling the impact of climate change on Mediterranean plant demography. [ABSTRACT FROM AUTHOR]

Destruction and fragmentation of habitats represent one of the most important threats for biodiversity. Here, we examined the effects of fragmentation in Mediterranean forests on the epiphytic lichen Lobaria pulmonaria (Lobariaceae). We tested the hypothesis that not only the level of fragmentation affects L. pulmonaria populations, but also the quality of the habitat and the nature of the surrounding matrix affect them. The presence and abundance of the lichen was recorded on 2039 trees in a total of 31 stands. We recorded habitat quality and landscape variables at three hierarchical levels: tree, plot, and patch. We found that L. pulinonaria tends to occur in trees with larger diameters in two types of surveyed forests. In Quercus pyrenaica patches, the mean diameter of colonized trees was smaller, suggesting the importance of bark roughness. Factors affecting the presence and cover of the lichen in each type of forest were different. There was a strong positive influence of distance from a river in beech forests, whereas proximity to forest edge positively affected in oak forests. The influence of the surrounding matrix was also an important factor explaining the epiphytic lichen abundance. [ABSTRACT FROM AUTHOR]

The worldwide phenomenon of shrub encroachment in grass-dominated dryland ecosystems is commonly associated with desertification. Studies of the purported desertification effects associated with shrub encroachment are often restricted to relatively few study areas, and document a narrow range of possible impacts upon biota and ecosystem processes. We conducted a study in degraded Mediterranean grasslands dominated by Stipa tenacissima to simultaneously evaluate the effects of shrub encroachment on the structure and composition of multiple biotic community components, and on various indicators of ecosystem function. Shrub encroachment enhanced vascular plant richness, biomass of fungi, actinomycetes and other bacteria, and was linked with greater soil fertility and N mineralization rates. While shrub encroachment may be a widespread phenomenon in drylands, an interpretation that this is an expression of desertification is not universal. Our results suggest that shrub establishment may be an important step in the reversal of desertification processes in the Mediterranean region. [ABSTRACT FROM AUTHOR]

Frequency-dependent processes are relevant for flowering plant reproduction, especially for species with disassortative mating. In an individual-based study, we tested not only the effects of local density on reproductive success at small spatial scales, but also those of neighborhood quality. To test the neighborhood effects on the reproduction of Armeria caespitosa, a dimorphic Mediterranean high-mountain endemic, we introduce a novel pollination context (PC) index that considered the distance, floral display, and floral morph of neighbors at small scales (within 2 m from the focal plant), studying rock and pasture populations at both edges of the species altitudinal distribution. Reproductive success depended significantly on PC only at the low populations, suggesting that the PC effects are population-dependent and supporting the hypothesis that the neighborhood quality influences the reproductive success of A. caespitosa at least in stressful conditions. Moreover, fruit set was morph-dependent in the high-pasture population. The specific role of the spatial structure of compatible vs. incompatible morphs at small scales of the dimorphic selfincompatibility system in Arineria deserves further attention. Parameters other than plant density are useful for the study of smallscale density-dependent processes that affect pollination and other reproductive components, especially if they integrate neighborhood quality information at adequate spatial scales. [ABSTRACT FROM AUTHOR]

According to life-history theory, reproductive investments involve costs in terms of growth, future fecundity, and/or survival. However, studies to date have often failed to detect costs of reproduction, with survival costs among the less documented. We investigated the cost of reproduction in Helianthernum squarnatuin (Cistaceae), a short-lived perennial of semiarid Mediterranean environments. After experimental flower removal, we evaluated next season's growth, reproduction, and survival of the plants. We also monitored an indicator of plant physiological status (Fv/Fm) and leaf nutrient concentration at key phenological stages during reproduction. Survival rate in deblossomed plants was significantly higher than in control plants. As far as we know, this is the first experimental evidence of a survival cost of reproduction in a perennial plant. In contrast, no cost to growth or reproduction was found during the next season, and no significant differences in Fv/Fm or leaf nutrients were found between control and deblossomed plants. Helianthemum squarnaturn's success in semiarid Mediterranean ecosystems seems to rely on a persistent seed bank, combined with a sustained high reproductive output at the expense of survival. We conclude that this strategy might be more common than previously thought among short-lived shrubby plants growing in stressful Mediterranean areas. [ABSTRACT FROM AUTHOR]

1. Summer drought stress is considered the primary constraint to plant performance in Mediterranean ecosystems. However, little is known about the implications of summer stress for plant reproduction under real field conditions and, particularly, for the regulatory mechanisms of maternal investment in reproduction. 2. The relationship between plant physiological status at different reproductive stages over the course of the summer drought period and final reproductive output was modelled in the Mediterranean semi-arid specialist Helianthemum squamatum. 3. Plant physiological status, assessed by the chlorophyll fluorescence-based parameter Fv/ Fm, and soil moisture content beneath each plant, were determined in the field at five key phenological moments in a total of 88 plants. We used Generalized Linear Mixed Models to evaluate the effect of plant physiological status at those different dates on several components of reproduction (number of flowers and seeds per plant, fruit-set and intra-fruit seed abortion). We included soil moisture as an additional predictor to statistically control its potential effect on reproduction. 4. Fv/ Fm measured at midday was a significant predictor of reproductive output, but its significance varied over time and with the specific reproductive response variable. Fv/ Fm measured at the onset of flowering was positively related to the number of flowers and seeds per plant, whereas Fv/ Fm at the fruiting peak positively affected fruit-set. Soil moisture content was only significant when measured before flowering, being positively related to total flowers and seeds. The effect of stress on reproductive output acted either at an early stage of the reproductive season, by varying the number of flowers produced and seed primordia initiated, or at a later stage, by adjusting the number or ripe fruits. 5. Synthesis. Our results show a direct relationship between physiological status and reproduction, and highlight the importance of the timing of stress for reproductive success. They also show that small departures from the physiological optimum at specific reproductive stages may cause significant decreases in the reproductive output. We suggest that the dynamic adjustment of reproduction in response to stress is adaptive in fluctuating and unpredictable Mediterranean semi-arid environments, where an adequate temporal distribution of maternal resources determines the species’ ability to withstand severe environmental conditions. [ABSTRACT FROM AUTHOR]

Vegetation in high Mediterranean mountains usually consists of patchy communities. Patch structures have been interpreted as a result of the prevalence of facilitation phenomena in highly stressful environments. Several mechanisms have been proposed in order to explain the factors that control the existence of these clumped structures. However, they have not been evaluated in these mountains. Our hypothesis is that patchy structure in high mountain Mediterranean vegetation is a consequence of facilitative and competitive interactions in a very harsh environment which ultimately involve strong localized effects on soil properties. Our results show that levels of soil nutrients were higher under vegetation patches than in bare ground areas, confirming the hypothesis of an amelioration of soil resources under canopies. Pairwise associations and repulsions suggest the existence of two contrasting composition stages. Contrasting models relating patch species composition (cover and biomass) and soil resources indicated a weak relationship between species features and soil nutrient levels. Finally, structural modeling showed that patch size has a relevant but indirect effect on soil resource levels through grass and total biomass. We conclude that patch structure and dynamics in high Mediterranean mountain communities may be partly controlled by an endogenous process involving facilitation and competition for soil key resources. These interactions may operate through some community traits related to patch size but not to composition. [ABSTRACT FROM AUTHOR]

Abstract: The co-occurrence pattern of epiphytic lichen communities was evaluated relative to micro-environmental heterogeneity along an edge-interior forest gradient. We collected data on the occurrence of 57 epiphytic lichen species from 452 plots in a Mediterranean forest remnant. We used two realistic null models to test for non-randomness in the structure of epiphytic lichen communities and generalized linear models to test the influence of micro-environmental variables on co-occurrence indices along the edge-interior gradient. We found that epiphytic lichen communities co-occurred less often than expected by chance, although the segregation pattern was not highly structured or recurrent. Less species co-occurrence was detected in the assemblage structure in the forest interior and with a southerly exposure. However, there were no significant relationships between co-occurrence and the number of coexisting species, the dominant species coverage or bryophyte coverage. In summary, our co-occurrence analyses suggested that epiphytic lichens were competitively structured, while we found no evidence of facilitation in more stressful conditions, as predicted by the stress-gradient hypothesis. Nevertheless, these results should be treated with caution because co-occurrence patterns may involve specific unknown modes of facilitation and competition, which relate to the life history of these organisms. [Copyright &y& Elsevier]

Abstract: Biological soil crusts (BSCs), composed of lichens, cyanobacteria, mosses, liverworts and microorganisms, are key biotic components of arid and semi-arid ecosystems worldwide. Despite they are widespread in Spain, these organisms have been historically understudied in this country. This trend is beginning to change as a recent wave of research has been identifying BSCs as a model ecological system. Many studies and research projects carried out in Spain have explored the role of BSCs on water, carbon and nitrogen fluxes, the interactions between BSCs and vascular plants, their dynamics after disturbances, and their response to global change, among other topics. In this article we review the growing body of research on BSCs available from semi-arid areas of Spain, highlighting its importance for increasing our knowledge on this group of organisms. We also discuss how it is breaking new ground in emerging research areas on the ecology of BSCs, and how it can be use to guide management and restoration efforts. Finally, we provide directions for future research on the ecology of BSCs in Spain and abroad. [Copyright &y& Elsevier]

Abstract: Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random. [Copyright &y& Elsevier]

Abstract: Shrub encroachment is a worldwide phenomenon with implications for desertification and global change. We evaluated its effects on the activities of urease, phosphatase and β-glucosidase in Mediterranean semiarid grasslands dominated by Stipa tenacissima by sampling 12 sites with and without resprouting shrubs along a climatic gradient. The presence of shrubs affected the evaluated enzymes at different spatial scales. Soils under S. tenacissima tussocks and in bare ground areas devoid of vascular plants had higher values of phosphatase and urease when the shrubs were present. For the β-glucosidase, this effect was site-specific. At the scale of whole plots (30 m × 30 m), shrubs increased soil enzyme activities between 2% (β-glucosidase) and 22% (urease), albeit these differences were significant only in the later case. Our results indicate that shrub encroachment does not reduce the activity of extracellular soil enzymes in S. tenacissima grasslands. [Copyright &y& Elsevier]

Background and Aims: Intra- and transgenerational plasticity may provide substantial phenotypic variation to cope with environmental change. Since assessing the unique contribution of the maternal environment to the offspring phenotype is challenging in perennial, outcrossing plants, little is known about the evolutionary and ecological implications of transgenerational plasticity and its persistence over the life cycle in these species. We evaluated how intra- and transgenerational plasticity interplay to shape the adaptive responses to drought in two perennial Mediterranean shrubs., Methods: We used a novel common garden approach that reduced within-family genetic variation in both the maternal and offspring generations by growing the same maternal individual in two contrasting watering environments, well-watered and drought, in consecutive years. We then assessed phenotypic differences at the reproductive stage between offspring reciprocally grown in the same environments., Key Results: Maternal drought had an effect on offspring performance only in Helianthemum squamatum. Offspring of drought-stressed plants showed more inflorescences, less sclerophyllous leaves and higher growth rates in both watering conditions, and heavier seeds under drought, than offspring of well-watered maternal plants. Maternal drought also induced similar plasticity patterns across maternal families, showing a general increase in seed mass in response to offspring drought, a pattern not observed in the offspring of well-watered plants. In contrast, both species expressed immediate adaptive plasticity, and the magnitude of intragenerational plasticity was larger than the transgenerational plastic responses., Conclusions: Our results highlight that adaptive effects associated with maternal drought can persist beyond the seedling stage and provide evidence of species-level variation in the expression of transgenerational plasticity. Such differences between co-occurring Mediterranean species in the prevalence of this form of non-genetic inheritance may result in differential vulnerability to climate change., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Although the role played by phylogeny in the assembly of plant communities remains as a priority to complete the theory of species coexistence, experimental evidence is lacking. It is still unclear to what extent phylogenetic diversity is a driver or a consequence of species assembly processes. We experimentally explored how phylogenetic diversity can drive the community level responses to drought conditions in annual plant communities. We manipulated the initial phylogenetic diversity of the assemblages and the water availability in a common garden experiment with two irrigation treatments: average natural rainfall and drought, formed with annual plant species of gypsum ecosystems of Central Spain. We recorded plant survival and the numbers of flowering and fruiting plants per species in each assemblage. GLMMs were performed for the proportion of surviving, flowering, fruiting plants per species and for total proportion of surviving species and plants per pot. In water limited conditions, high phylogenetic diversity favored species coexistence over time with higher plant survival and more flowering and fruiting plants per species and more species and plants surviving per pot. Our results agree with the existence of niche complementarity and the convergence of water economy strategies as major mechanisms for promoting species coexistence in plant assemblages in semiarid Mediterranean habitats. Our findings point to high phylogenetic diversity among neighboring plants as a plausible feature underpinning the coexistence of species, because the success of each species in terms of surviving and producing offspring in drought conditions was greater when the initial phylogenetic diversity was higher. Our study is a step forward to understand how phylogenetic relatedness is connected to the mechanisms determining the maintenance of biodiversity., (© 2021. The Author(s).)

Techniques for retrospective analysis of size dynamics at annual resolution remain poorly developed in lichens in general, and fruticose lichens in particular. Only a few attempts in very high latitudes suggested that growth might be studied as a chronosequence of inter-nodal branch elongations. Here we evaluated, for the first time, this hypothesis in a dry Mediterranean environment using the lichen Cladonia rangiformis as a case study. Mixed models supported a strong positive relationship between humidity measured as precipitation/PET and inter-nodal branch elongations. Importantly, model selection suggested that (i) the number of intermodal elongations were a major determinant of stem elongation, and (ii) a second-order temporal autocorrelation denoted legacies of environmental influences at least over the next 2 y. The strong growth-humidity relationship, along with the potential legacies observed, support the idea that inter-nodal branch elongations could be used to reconstruct growth chronologies at annual resolution in drylands. This finding highlights the high vulnerability of these organisms to rising aridity, and opens a new venue for climate reconstruction and other potential applications in Ecology and Earth Science disciplines., (Copyright © 2019 British Mycological Society. Published by Elsevier Ltd. All rights reserved.)

High-mountain areas provide excellent opportunities to study the effects of combined abiotic stresses on plant physiology given their variety of steep ecological gradients, low anthropogenic disturbance and remarkable levels of taxonomic diversity. Efficient photoprotective and antioxidant scavenging mechanisms are vital for survival in high-mountain plants, having its altitudinal and seasonal variations determined by environmental or ontogenetic factors such as the decrease in mean temperatures and water availability. A number of stress indicators have been described in order to rapidly assess plant fitness in high-mountain environments. For instance, carbon isotope (δ 13 C) and proline content as drought and temperature stress indicators, because of their link to water-use efficiency and osmotic adjustment; photosynthetic pigments, related to phenology, nutrient status, light and temperature stress; and non-structural carbohydrate accumulation in response to mild or brief drought conditions. The present review unveils the wide research opportunities available for the study of adaptive responses in high-mountain plants via stress indicators, and calls attention to the substantial knowledge gap existing between alpine zones and other mountainous regions, such as Mediterranean high-mountains. The aim is to grant a more holistic understanding of the physiological mechanisms driving plant life in high altitudes and improve the predictions of the effects of changing environments in these species and across ecological scales., (© 2018 Scandinavian Plant Physiology Society.)

Two-phase plant communities with an engineer conforming conspicuous patches and affecting the performance and patterns of coexisting species are the norm under stressful conditions. To unveil the mechanisms governing coexistence in these communities at multiple spatial scales, we have developed a new point-raster approach of spatial pattern analysis, which was applied to a Mediterranean high mountain grassland to show how Festuca curvifolia patches affect the local distribution of coexisting species. We recorded 22 111 individuals of 17 plant perennial species. Most coexisting species were negatively associated with F. curvifolia clumps. Nevertheless, bivariate nearest-neighbor analyses revealed that the majority of coexisting species were confined at relatively short distances from F. curvifolia borders (between 0-2 cm and up to 8 cm in some cases). Our study suggests the existence of a fine-scale effect of F. curvifolia for most species promoting coexistence through a mechanism we call 'facilitation in the halo'. Most coexisting species are displaced to an interphase area between patches, where two opposite forces reach equilibrium: attenuated severe conditions by proximity to the F. curvifolia canopy (nutrient-rich islands) and competitive exclusion mitigated by avoiding direct contact with F. curvifolia., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

Climate change is imposing warmer and more arid conditions on high-mountain Mediterranean pastures. The severity of these conditions is more intense in lower elevation populations and may be critical for their survival. In this context, we asked whether local adaptation plays an important role in the response of these populations to climate change, and if so, what mechanisms are involved. Previous works, involving reciprocal sowings suggested the existence of local adaptation in lower elevation populations of Silene ciliata, a perennial representative of high-mountain Mediterranean pastures. To determine if this local advantage is due to better adaptation to more intense water stress conditions, an experiment was conducted in which S. ciliata plants from three populations located at different elevations (Low, Intermediate and High) were subjected to severe artificial water stress. Results showed that plants from the Low population had greater tolerance to water stress than plants from the High population in the earliest stages of water shortage. Furthermore, responses of proteins to specific antibodies related to drought were evaluated. Two representative late-embryogenesis abundant (LEA) proteins known to play a role in water stress tolerance were expressed throughout the drought treatment in plants from the three populations, with some pattern differences among individuals within populations. This study detected slight evidence of local adaptation to water stress in populations from different elevations., (© 2012 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Global change triggers shifts in forest composition, with warming and aridification being particularly threatening for the populations located at the rear edge of the species distributions. This is the case of Scots pine (Pinus sylvestris) in the Mediterranean Basin where uncertainties in relation to its dynamics under these changing scenarios are still high. We analysed the relative effect of climate on the recruitment patterns of Scots pine and its interactions with local biotic and abiotic variables at different spatial scales. Number of seedlings and saplings was surveyed, and their annual shoot growth measured in 96 plots located across altitudinal gradients in three different regions in the Iberian Peninsula. We found a significant influence of climate on demography and performance of recruits, with a non-linear effect of temperature on the presence of juveniles, and a positive effect of precipitation on their survival. Abundance of juveniles of P. sylvestris that underwent their first summer drought was skewed towards higher altitudes than the altitudinal mean range of the conspecific adults and the optimum elevation for seedlings' emergence. At local level, light availability did not influence juveniles' density, but it enhanced their growth. Biotic interactions were found between juveniles and the herb cover (competition) and between the number of newly emerged seedlings and shrubs (facilitation). Results also highlighted the indirect effect that climate exerts over the local factors, modulating the interactions with the pre-existing vegetation that were more evident at more stressful sites. This multiscale approach improves our understanding of the dynamics of these marginal populations and some management criteria can be inferred to boost their conservation under the current global warming.

Background and Aims: In mountain plant populations, local adaptation has been described as one of the main responses to climate warming, allowing plants to persist under stressful conditions. This is especially the case for marginal populations at their lowest elevation, as they are highly vulnerable. Adequate levels of genetic diversity are required for selection to take place, while high levels of altitudinal gene flow are seen as a major limiting factor potentially precluding local adaptation processes. Thus, a compromise between genetic diversity and gene flow seems necessary to guarantee persistence under oncoming conditions. It is therefore critical to determine if gene flow occurs preferentially between mountains at similar altitudinal belts, promoting local adaptation at the lowest populations, or conversely along altitude within each mountain., Methods: Microsatellite markers were used to unravel genetic diversity and population structure, inbreeding and gene flow of populations at two nearby altitudinal gradients of Silene ciliata, a Mediterranean high-mountain cushion plant., Key Results: Genetic diversity and inbreeding coefficients were similar in all populations. Substantial gene flow was found both along altitudinal gradients and horizontally within each elevation belt, although greater values were obtained along altitudinal gradients. Gene flow may be responsible for the homogeneous levels of genetic diversity found among populations. Bayesian cluster analyses also suggested that shifts along altitudinal gradients are the most plausible scenario., Conclusions: Past population shifts associated with glaciations and interglacial periods in temperate mountains may partially explain current distributions of genetic diversity and population structure. In spite of the predominance of gene flow along the altitudinal gradients, local genetic differentiation of one of the lower populations together with the detection of one outlier locus might support the existence of different selection forces at low altitudes.

Reproduction at population lower edges is important for plant species persistence, especially in populations on contracting high-mountain islands. In this context, the ability of plants to reproduce in different microhabitats seems to be important to guarantee seed production in stressful environments, such as Mediterranean high mountains. We hypothesised that the warmer and drier conditions at the lower edge would be deleterious for the reproduction of Armeria caespitosa, an early-flowering plant. In addition, reproductive plasticity along this mountain gradient may also be microhabitat-dependent. We studied factors affecting the reproductive success of A. caespitosa, an endemic of the Spanish Sistema Central. We considered a complex set of predictors, including phenology, plant size and environmental factors at different scales using generalised estimating equations and generalised linear models. Microhabitat, together with the position in the altitudinal gradient and inter-annual variability affected the reproduction of A. caespitosa. In addition, individuals with longer flowering periods (duration of flowering) had significantly lower fruit set and a higher number of unviable seeds; delayed flowering peaks favoured the production of both viable and unviable fruits. Microhabitat variability over an altitudinal range is relevant for the reproduction of A. caespitosa, and is more important at the lower edge of the altitudinal range, where the species faces the most adverse conditions. In addition, the ability to reproduce in different microhabitats might increase the chances of the species to cope with environmental uncertainties under on-going climate warming. Finally, reproduction of this early-flowering plant is constrained by summer drought, which might shape its reproductive phenology.

Mountain plants are particularly sensitive to climate warming because snowmelt timing exerts a direct control on their reproduction. Current warming is leading to earlier snowmelt dates and longer snow-free periods. Our hypothesis is that high-mountain Mediterranean plants are not able to take advantage of a lengthened snow-free period because this leads to longer drought that truncates the growing season. However, reproductive timing may somewhat mitigate these negative effects through temporal shifts. We assessed the effects of flowering phenology on the reproductive success of Silene ciliata, a Mediterranean high-mountain plant, across an altitudinal gradient during two climatically contrasting years. The species showed a late-flowering pattern hampering the use of snowmelt water. Plant fitness was largely explained by the elapsed time from snowmelt to onset of flowering, suggesting a selective pressure towards early flowering caused by soil moisture depletion. The proportion of flowering plants decreased at the lowest population, especially in the drier year. Plants produced more flowers, fruits and seeds at the highest population and in the mild year. Our results indicate that water deficit in dry years could threaten the lowland populations of this mountainous species, while high-altitude environments are more stable over time.

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.

Nitrogen uptake, nitrogen demand and internal nitrogen cycling were studied to address the question of the importance of nutrient storage in Quercus species with contrasting leaf longevities. We carried out this study at the whole-plant level with young trees (2-4 years old) of three Mediterranean Quercus species: the evergreen Q. ilex, the marcescent/evergreen Q. faginea, and the deciduous Q. pyrenaica. Seasonal dynamics of nitrogen in all compartments of the plant were followed for 3 years. Nitrogen losses were measured through litter production, herbivory and fine root shedding. Nitrogen uptake was estimated using increments of nitrogen plant content plus accumulative nitrogen losses. Nitrogen uptake was limited to a few months during late winter and spring. Before budbreak, acquired nitrogen was stored in old-leaf cohorts of evergreen and woody compartments. After budbreak, Quercus species relied first on soil uptake and second on nitrogen retranslocation to supply new growth requirements. However, in most cases we found a high asynchrony between nitrogen demand by growing tissues and soil supply, which determined a strong nitrogen retranslocation up to 88.4% of the nitrogen demand throughout leaf expansion. Except for the first year after planting, the above- and underground woody fractions provided more nitrogen to the new tissues than the old leaf cohorts. Differences in the benefit of nitrogen withdrawn from senescent and old leaves were not found between species. We conclude that sink/source interaction strength was determined by differences between nitrogen demand and uptake, regulating internal nutrient cycling at the whole plant level.