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Eucalyptus globulus Labill. is one of the most widely planted hardwood species worldwide. This species occupies a quarter of the country's forested area in Portugal, so there is a growing concern about its post-fire dispersal. Although it is generally recognised that fire promotes E. globulus natural regeneration and that precipitation and topography influence recruitment, little is known about the role of post-fire conditions on the establishment of the species following off-season fires. We examine how post-fire conditions affect E. globulus natural regeneration and invasive potential. Sapling establishment was assessed in plantations and under old and large isolated eucalyptus trees (seed-trees) following two off-season fire events (2017 June and October fires). Abiotic and biotic local factors affected differently sapling establishment in plantations and under seed-trees. In plantations, sapling cover was more influenced by tree-related traits (age and height), whereas the conditions beneath seed-trees mitigated the impact of harsh conditions on saplings. In both cases, post-fire climatic conditions constrained natural regeneration, with less impact under seed-trees, reinforcing their importance in establishment success. Thus, isolated seed-trees should be considered while managing the species' unplanned spread. [ABSTRACT FROM AUTHOR]

Aims: Our aim was to investigate the water sources used by woody species under contrasting water availability and the extent of water-sources-use differentiation among dominant woody species in a tropical coastal dune forest. Methods: We sampled 15 woody species in a Brazilian restinga forest and, through Bayesian isotope mixing models, we estimated the proportion of water sources used. We tested whether water-sources-use was (i) different between contrasting water availability conditions; (ii) dependent on growth form, plant size or crown illumination; and (iii) influenced by stand density, evenness or biomass. Results: We found a seasonal variation in water-sources-use, but no vertical soil-water partitioning among woody species. In wetter periods, plants used mainly water from top-soil, as a shallow water table limited water uptake to top-soil layers recharged with rainwater. Contrastingly, during drier periods, with the absence of rain and a deeper water table, plants generally relied on deeper (50 cm) soil layers. Only under less-wet conditions, a greater evenness and density implied higher water-uptake depth heterogeneity among plants. However, changes in the main water-sources used by plants were neither evoked in more dense or diverse plots, nor induced by plant size. Conclusions: Our study shows that restinga species have dynamic shifts in water-uptake depth caused by seasonal water availability changes, influenced by the combined effect of insufficient moisture at shallow soil layers and water-table lowering in drier periods. These temporal shifts are common among species, implying that restinga woody community has a homogeneous strategy of water-resources acquisition. This study enhances our understanding of the effects that water variations can have on water-resource use in restinga forests. [ABSTRACT FROM AUTHOR]

The pinewood nematode, Bursaphelenchus xylophilus, originating from North America ( NA), is a major invasive pine pest in Eurasia. It was first detected in Portugal in 1999 associated with maritime pine, Pinus pinaster, and has been differently affecting the main local pine species, P. pinaster and P. pinea. Field studies and direct inoculation experiments in Pinus spp. seedlings, under controlled conditions, were performed to assess whether the differences in constitutive and inducible defences are determining the different susceptibility of pine host species to B. xylophilus. Host co-evolution with the pathogen was also assessed, including the NA P. radiata, widely used in forestry in the northeast of the Iberian peninsula. Pine mortality in the field was positively related with the abundance of B. xylophilus, and concentration of phenolics and condensed tannins in pines. In the greenhouse assay, seedling tissues were analysed for constitutive investment in defences, as well as the potential inducibility of those defences as driven by B. xylophilus inoculation. Slower growing P. pinea presented higher levels of constitutive defences than faster growing P. pinaster, with only P. pinaster being affected by B. xylophilus. Furthermore, co-evolution with the pathogen is important, with the fast-growing NA P. radiata presenting an inducible and effective response to B. xylophilus. Results point to the importance of integrating data on pine life history traits, including growth rate, and production of constitutive and inducible defences, into predictive models for this invasive forest pest. [ABSTRACT FROM AUTHOR]

Stable isotope analysis is a powerful tool for assessing plant carbon and water relations and their impact on biogeochemical processes at different scales. Our process-based understanding of stable isotope signals, as well as technological developments, has progressed significantly, opening new frontiers in ecological and interdisciplinary research. This has promoted the broad utilisation of carbon, oxygen and hydrogen isotope applications to gain insight into plant carbon and water cycling and their interaction with the atmosphere and pedosphere. Here, we highlight specific areas of recent progress and new research challenges in plant carbon and water relations, using selected examples covering scales from the leaf to the regional scale. Further, we discuss strengths and limitations of recent technological developments and approaches and highlight new opportunities arising from unprecedented temporal and spatial resolution of stable isotope measurements. [ABSTRACT FROM AUTHOR]

Nitrogen (N) has emerged in recent years as a key factor associated with global changes, with impacts on biodiversity, ecosystems functioning and human health. In order to ameliorate the effects of excessive N, safety thresholds such as critical loads (deposition fluxes) and levels (concentrations) can be established. Few studies have assessed these thresholds for semi-natural Mediterranean ecosystems. Our objective was therefore to determine the critical loads of N deposition and long-term critical levels of atmospheric ammonia for semi-natural Mediterranean evergreen woodlands. We have considered changes in epiphytic lichen communities, one of the most sensitive community indicators of excessive N in the atmosphere. Based on a classification of lichen species according to their tolerance to N we grouped species into response functional groups, which we used as a tool to determine the critical loads and levels. This was done for a Mediterranean climate in evergreen cork-oak woodlands, based on the relation between lichen functional diversity and modelled N deposition for critical loads and measured annual atmospheric ammonia concentrations for critical levels, evaluated downwind from a reduced N source (a cattle barn). Modelling the highly significant relationship between lichen functional groups and annual atmospheric ammonia concentration showed the critical level to be below 1.9 µgm-3, in agreement with recent studies for other ecosystems. Modelling the highly significant relationship between lichen functional groups and N deposition showed that the critical load was lower than 26 kg (N) ha-1 yr-1, which is within the upper range established for other semi-natural ecosystems. Taking into account the high sensitivity of lichen communities to excessive N, these values should aid development of policies to protect Mediterranean woodlands from the initial effects of excessive N. [ABSTRACT FROM AUTHOR]

In spite of the relative importance of groundwater in costal dune systems, studies concerning the responses of vegetation to ground water (GW) availability variations, particularly in Mediterranean regions, are scarce. Thus, the main purpose of this study is to compare the responses of co-occurring species possessing different functional traits, to changes in GW levels (i.e. the lowering of GW levels) in a sand dune ecosystem. For that, five sites were established within a 1 km² area in a meso-mediterranean sand dune ecosystem dominated by a Pinus pinaster forest. Due to natural topographic variability and anthropogenic GW exploitation, substantial variability in depth to GW between sites was found. Under these conditions it was possible to identify the degree of usage and dependence on GW of different plant species (two deep-rooted trees, a drought adapted shrub, a phreatophyte and a non-native woody invader) and how GW dependence varied seasonally and between the heterogeneous sites. Results indicated that the plant species had differential responses to changes in GW depth according to specific functional traits (i.e. rooting depth, leaf morphology, and water use strategy). Species comparison revealed that variability in pre-dawn water potential (ψpre) and bulk leaf δ13C was related to site differences in GW use in the deeprooted (Pinus pinaster, Myrica faya) and phreatophyte (Salix repens) species. However, such variation was more evident during spring than during summer drought. The exotic invader, Acacia longifolia, which does not possess a very deep root system, presented the largest seasonal variability in ψpre and bulk leaf δ13C. In contrast, the response of Corema album, an endemic understory drought-adapted shrub, seemed to be independent of water availability across seasons and sites. Thus, the susceptibility to lowering of GW due to anthropogenic exploitation, in plant species from sand dunes, is variable, being particularly relevant for deep rooted species and phreatophytes, which seem to depend heavily on access to GW. [ABSTRACT FROM AUTHOR]

Stable isotope analysis is a powerful tool for tracing biogeochemical processes in the carbon and water cycles. One particularly powerful approach is to employ multiple isotopes where the simultaneous assessment of the D/H,18O/16O and/or 13C/12C in different compounds provide a unique means to investigate the coupling of water and 5 carbon fluxes at various temporal and spatial scales. Here, we present a research update on recent advances in our process-based understanding of the utilization of carbon, oxygen and hydrogen isotopes to lend insight into carbon and water cycling. We highlight recent technological developments and approaches, their strengths and methodological precautions with examples covering scales from minutes to centuries and from the leaf to the globe. [ABSTRACT FROM AUTHOR]

In spite of the relative importance of groundwater in coastal dune systems, the number of studies concerning the responsiveness of vegetation to ground water (GW) variability, in particularly in Mediterranean regions, is scarce. In this study, we established 5 study sites within a meso-mediterranean sand dune Pinus pinaster forest on the Atlantic coast of Portugal, taking advantage of natural topographic variability and artificial GW exploitation, which resulted in substantial variability in depth to GW between microsites. Here we identify the degree of usage and dependence on GW of different plant functional groups (two deep-rooted trees, a drought adapted shrub, a phreatophyte and a non-native woody invader) and how GW dependence varied seasonally and between the heterogeneous microsites. Our results indicated that the plant species had differential responses to changes in GW depth according to specific functional traits (i.e. rooting depth, leaf morphology, and water use strategy). The species comparison revealed that variability in pre-dawn water potential (ψpre) and bulk leaf δ 13C was related to microsite differences in GW use in deep-rooted (Pinus pinaster, Myrica faya) and phreatophyte (Salix repens) species. However, such variation was more evident during spring rather than during summer drought. The exotic invader, Acacia longifolia, which does not possess a very deep root system, presented the largest seasonal variability in ψpre and bulk leaf δ 13C. In contrast, the response of Corema album, an endemic understorey drought adapted shrub, across seasons and microsites seemed to be independent of water availability. Thus, the susceptibility to changing GW availability in sand dune plant species is variable, being particularly relevant for deep rooted species and phreatophytes, which have typically been less exposed to GW fluctuations. [ABSTRACT FROM AUTHOR]

The study presents first data on the δ18O performance of poikilohydrous lichen ground cover, and its potential impact on the isotopic composition of water fluxes arising from subjacent soil layers. As a model organism, the globally distributed lichen Cladina arbuscula was studied under laboratory conditions as well as in the field. During a desiccation experiment, δ18O of the lichen's thallus water and of its respired CO2 became enriched by ∼7‰ and followed a similar enrichment pattern to that expected from homoiohydrous, vascular plants. However, the observed degree of enrichment was lower in comparison to vascular plants due to (i) the lichen's inherent lower evaporative resistances; and (ii) a stronger effect of the more depleted surrounding water vapour. In lichens growing in their natural habitat, this specific pattern may show substantial variations depending on prevailing microclimatic conditions. Within a field study, thallus water δ18O of lichens principally proved to become more depleted when close to equilibration with the surroundings. It thereby strongly depended on the absorption of surrounding water vapour. Moreover, the results indicate that lichen mats substantially reduce evaporation rates arising from subjacent soil layers, and may alter the isotopic signal of vapour diffusing away from these layers into more depleted values. [ABSTRACT FROM AUTHOR]

Limestone quarries are spread over the Mediterranean Basin and have a strong environmental impact on the landscape, causing vegetation losses and soil losses. A reclamation project was conducted in a limestone quarry, situated in Arrábida Natural Park (southwest Portugal), that is dominated by mediterranean vegetation. Revegetation was conducted using three evergreen sclerophyllous shrub species (Ceratonia siliqua, Olea europaea, and Pistacia lentiscus), and new techniques were assayed to improve plant water status and nutrient status during the first phases after plantation. A water-holding polymer (gel), fertilizer, and mycorrhiza inoculum were applied in a factorial experiment in a randomized complete block design. The success of these techniques was evaluated during 1.5 years, through monitoring of growth and ecophysiology of plants. Plant survival was high, the lowest values (95%) being recorded in C. siliqua. There were species-specific responses to the treatments applied. Ceratonia siliqua showed the highest growth rates and was the only species with growth stimulated by fertilizer application. However, the application of fertilizer induced changes in leaf characteristics of the other two species, increasing chlorophyll and nitrogen contents. Mycorrhiza inoculum had no effect on plant response. The addition of the water-holding polymer induced higher midday plant water potentials in C. siliqua, O. europaea, and P. lentiscus, but in the latter two species the simultaneous addition of gel and fertilizer induced the lowest water potentials. The addition of fertilizer and gel is recommended in future revegetation programs but not the combination of both when revegetating with O. europaea and P. lentiscus. The results of this experiment indicate that the use of these mediterranean species, adapted to nutrient and water stress, can circumvent harsh conditions of the quarry. [ABSTRACT FROM AUTHOR]

The characteristics of gas exchange and carbon isotope discrimination were determined for a number of lichen species, representing contrasting associations between fungal (mycobiont) and photosynthetic (photobiont) organism. These parameters were evaluated with regard to the occurrence of any CO-concentrating mechanism (CCM) expressed specifically by the green algal (phycobiont) or cyanobacterial (cyanobiont) partner. Carbon isotope discrimination (Δ) fell into three categories. The highest Δ, found in lichens comprising a phycobiont plus cyanobacteria limited to pockets in the thallus (known as cephalodia), ranged from 24 to 28‰, equivalent to a carbon isotope ratio (δC) of around -32 to-36‰ vs. Pee Dee Belemnite (PDB) standard. Further evidence was consistent with CO supply to the carboxylating system entirely mediated by diffusion rather than a CCM, in that thallus CO compensation point and online instantaneous Δ were also high, in the range normally associated with C higher plants. For lichens consisting of phycobiont or cyanobiont alone, organic material Δ formed two distinct ranges around 15‰ (equivalent to a δC of -23%.). Thallus compensation point and instantaneous Δ were lower in the cyanobiont group, which also showed higher maximum rates of net photosynthesis, whether expressed on the basis of thallus dry weight, chlorophyll content or area. These data provide additional evidence for the activity of a CCM in cyanobiont lichens, which only show photosynthetic activity when reactivated with liquid water. Rates of net CO uptake were lower in both phycobiont associations, but were relatively constant across a wide working range of thallus water contents, usually in parallel with on-line Δ. The phycobiont response was consistent whether photosynthesis had been reactivated with liquid water or water vapour. The effect of diffusion limitation could generally be seen with a 3-4‰ decrease in instantaneous Δ at the highest water contents. The expression of a CCM in phycobiont algae, although reduced compared with that in cyanobacteria, has already been related to the occurrence of pyrenoids in chloroplasts. In view of the inherent requirement of cyanobacteria for some form of CCM, and the smaller pools of dissolved inorganic carbon (DIC = CO + HCO+ CO) associated with phycobiont lichens, it appears that Δ characteristics provide a good measure of the magnitude of any CCM, albeit tempered by diffusion limitation at the highest thallus water contents. [ABSTRACT FROM AUTHOR]

Variations in stable carbon isotope discrimination (Δ) were investigated across the thalli of several lichen species possessing different photobiont associations. Lichens containing (i) green algae (phycobiont), (ii) green algae in association with cyanobacteria confined in cephalodia, or (iii) cyanobacteria (cyanobiont) as the photobiont partner were studied. Carbon isotope discrimination was analysed in different thallus sections, which varied in distance from the margin and in age. The marginal thallus region is considered to be youngest, while the central region is thought to be oldest. This analysis showed a clear variation in Δ across the thallus related to distance from the growing margin. In most of the species examined, the highest Δ values were found in marginal regions (younger), while the central and basal regions (older) showed significantly lower Δ. To investigate the effects of the historical increase in atmospheric CO2 concentration and the concurrent decrease in the 13C content of atmospheric CO2 on the Δ of lichens, experiments were carried out on herbarium samples of tobaria pulmonaria collected from the mid 19th Century to 1953. The results obtained showed a pattern of variation of Δ consistent with that of freshly collected samples; A decreased substantially with increasing distance from the thallus margin, irrespective of the collection date. Moreover, no consistent variation of discrimination was found among different collection dates. These results demonstrate that the observed variation in Δ is caused by age-related changes in the physiological behaviour of different thallus sections, and that the past 150 years of increasing CO concentration have not had significant effects on Δ in L. pulmonaria. Photosynthesis measurements, chlorophyll analysis and observations using optical microscopy, performed on freshly collected lichens, showed significant changes in morphological and physiological characteristics across the thallus. Particularly, remarkable variations in thickness were found across the thallus. These anatomical changes may be responsible for the variation in Δ, through variations in CO2 transfer resistance and, consequently, in CO2 availability across the thallus. The lack of age-dependent variation in Δ in cyanobiont lichens is possibly attributable to the operation of a CO2 -concentrating mechanism and, therefore, to a more constant CO2 environment across the thallus in this lichen group. [ABSTRACT FROM AUTHOR]