Identification of potential invasive alien species in Spain through horizon scanning.

Invasive alien species have widespread impacts on native biodiversity and ecosystem services. Since the number of introductions worldwide is continuously rising, it is essential to prevent the entry, establishment and spread of new alien species through a systematic examination of future potential threats. Applying a three-step horizon scanning consensus method, we evaluated non-established alien species that could potentially arrive, establish and cause major ecological impact in Spain within the next 10 years. Overall, we identified 47 species with a very high risk (e.g. Oreochromis niloticus, Popillia japonica, Hemidactylus frenatus, Crassula helmsii or Halophila stipulacea), 61 with high risk, 93 with moderate risk, and 732 species with low risk. Many of the species categorized as very high or high risk to Spanish biodiversity are either already present in Europe and neighbouring countries or have a long invasive history elsewhere. This study provides an updated list of potential invasive alien species useful for prioritizing efforts and resources against their introduction. Compared to previous horizon scanning exercises in Spain, the current study screens potential invaders from a wider range of terrestrial, freshwater, and marine organisms, and can serve as a basis for more comprehensive risk analyses to improve management and increase the efficiency of the early warning and rapid response framework for invasive alien species. We also stress the usefulness of measuring agreement and consistency as two different properties of the reliability of expert scores, in order to more easily elaborate consensus ranked lists of potential invasive alien species.

Quantifying carbon storage and sequestration by native and non-native forests under contrasting climate types.

Non-native trees may have significant impacts on the carbon sink capacity of forested lands. However, large-scale patterns of the relative capacity of native and non-native forests to uptake and store carbon remain poorly described in the literature, and this information is urgently needed to support management decisions. In this study, we analyzed 17,065 plots from the Spanish Forest Inventory (covering c. 30 years) to quantify carbon storage and sequestration of natural forests and plantations of native and non-native trees under contrasting climate types, while controlling for the effects of environmental factors (forest structure, climate, soil, topography, and management). We found that forest origin (non-native vs. native) highly influenced carbon storage and sequestration, but such effect was dependent on climate. Carbon storage was greater in non-native than in native forests in both wet and dry climates. Non-native forests also had greater carbon sequestration than native ones in the wet climate, due to higher carbon gains by tree growth. However, in the dry climate, native forests had greater carbon gains by tree ingrowth and lower carbon loss by tree mortality than non-native ones. Furthermore, forest type (classified by the dominant species) and natural forests versus tree plantations were important determinants of carbon storage and sequestration. Native and non-native Pinus spp. forests had low carbon storage, whereas non-native Eucalyptus spp. forests and native Quercus spp., Fagus sylvatica, and Eurosiberian mixed forests (especially not planted ones) had high carbon storage. Carbon sequestration was greatest in Eucalyptus globulus, Quercus ilex, and Pinus pinaster forests. Overall, our findings suggest that the relative capacity of native and non-native forests to uptake and store carbon depends on climate, and that the superiority of non-native forests over native ones in terms of carbon sequestration declines as the abiotic filters become stronger (i.e., lower water availability and higher climate seasonality).

Searching for predictors of the variability of impacts caused by non-native trees on regulating ecosystem services worldwide.

Humans have introduced non-native trees (NNT) all over the world to take advantage of the plethora of benefits they provide. However, depending on the context, NNT may present a diverse range of effects on ecosystem services (ES), from benefits to drawbacks, which may hinder the development of policies for these species. Unfortunately, the attempts so far to understand the impacts of NNT on ES only explained a low proportion of their variation. Here we analyze the variation in impacts of NNT on regulating ecosystem services (RES) by using a global database, which covers the effect size of multiple NNT species on six RES (climate regulation, soil erosion regulation, soil fertility, soil formation, hydrological cycle regulation, and fire protection). We used a wide range of predictors to account for the context-dependency of impacts distributed in five groups: the RES type, functional traits of both the NNT and the dominant NT of the recipient ecosystem, phylogenetic and functional distances between NNT and NT, climatic context, and human population characteristics. Using boosted regression trees and regression trees, we found that the most influential predictors of NNT impacts on RES were annual mean temperatures and precipitation seasonality, followed by the type of RES, human population density, and NNT height. In regions with warm temperatures and low seasonality, NNT tended to increase RES. NNT impacts were greater in densely populated regions. Smaller NNT exerted greater positive impacts on climate regulation and soil erosion regulation in tropical regions than in other climates. We highlight that benign climates and high population density exacerbate the effects of NNT on RES, and that soil fertility is the most consistently affected RES. Knowledge of the factors that modulate NNT impacts can help to predict their potential effects on RES in different parts of the world and at various environmental settings.

Effects of widespread non-native trees on regulating ecosystem services.

Tree taxa are often planted beyond their native range to increase the provision of some ecosystem services. Yet, they can disrupt ecosystem processes in their new ranges, causing changes in the provision of other services. Here we review the effects of five widespread tree taxa (Acacia, Ailanthus, Eucalyptus, Pinus and Robinia) on six regulating ecosystem services in areas where they are non-native. We conducted a literature search for pair-wise comparisons between sites dominated by any of the selected taxa and sites with native vegetation. An array of variables were used as indicators for each ecosystem service. Data were analysed using multi-level meta-analyses to compare effects of taxa on each ecosystem service, and effects of the same taxa across contexts. We compiled 857 case studies from 107 source papers. Several taxa tended to increase climate regulation, mostly Eucalyptus. Acacia decreased fire risk prevention. Robinia, Acacia and Ailanthus increased soil fertility, while Eucalyptus and Pinus, tended to decrease it. Soil formation was enhanced by Robinia and Ailanthus. Acacia promoted the increase of water in land pools, while Eucalyptus tended to decrease them. All effects show a large heterogeneity across case studies. Part of this heterogeneity could be attributed to gross climatic differences (i.e. biome), to species differences within each genus, to the structure of the recipient ecosystem, and/or to human management. Managers and policy-makers should consider the context-dependency and the potential effects of non-native trees on a wide range of services to ground their decisions. Our analyses also revealed important gaps of knowledge (e.g. on fire risk prevention, erosion control or water cycle regulation) and some potential publication bias. The methodology used here easily allows for future updates as new information will become available.

A Global Review of Ligustrum Lucidum (OLEACEAE) Invasion.

Ligustrum lucidum is a highly invasive East Asian tree that successfully colonizes several subtropical and temperate areas around the world. Its invasion capacity results from a widespread human use mostly in urban and periurban settings, very abundant fruit and seed production, small bird-dispersed fruits, high germination rates, resprouting capacity, fast growth rates, low herbivory levels and tolerance to a wide range of light, temperature and soil. All these traits contribute to its ability to rapidly increase in abundance, alter biodiversity, landscape ecology and limit its management. This paper reviews the current knowledge on L. lucidum with particular focus on its uses, distribution, invasiveness, ecological and economic impacts and control measures. Most relevant aspect of the review highlight the negative ecological impacts of L. lucidum, its potential to continue expanding its range of distribution and the need of further studies on the eco-physiology of the species, economic impact and social perception of its invasion and early warning systems.

Allometric co-variation of xylem and stomata across diverse woody seedlings.

Leaf stomatal density is known to co-vary with leaf vein density. However, the functional underpinning of this relation, and how it scales to whole-plant water transport anatomy, is still unresolved. We hypothesized that the balance of water exchange between the vapour phase (in stomata) and liquid phase (in vessels) depends on the consistent scaling between the summed stomatal areas and xylem cross-sectional areas, both at the whole-plant and single-leaf level. This predicted size co-variation should be driven by the co-variation of numbers of stomata and terminal vessels. We examined the relationships of stomatal traits and xylem anatomical traits from the entire plant to individual leaves across seedlings of 53 European woody angiosperm species. There was strong and convergent scaling between total stomatal area and stem xylem area per plant and between leaf total stomatal area and midvein xylem area per leaf across all the species, irrespective of variation in leaf habit, growth-form or relative growth rate. Moreover, strong scaling was found between stomatal number and terminal vessel number, whereas not in their respective average areas. Our findings have broad implications for integrating xylem architecture and stomatal distribution and deepen our understanding of the design rules of plants' water transport network.

Global effects of non-native tree species on multiple ecosystem services.

Non-native tree (NNT) species have been transported worldwide to create or enhance services that are fundamental for human well-being, such as timber provision, erosion control or ornamental value; yet NNTs can also produce undesired effects, such as fire proneness or pollen allergenicity. Despite the variety of effects that NNTs have on multiple ecosystem services, a global quantitative assessment of their costs and benefits is still lacking. Such information is critical for decision-making, management and sustainable exploitation of NNTs. We present here a global assessment of NNT effects on the three main categories of ecosystem services, including regulating (RES), provisioning (PES) and cultural services (CES), and on an ecosystem disservice (EDS), i.e. pollen allergenicity. By searching the scientific literature, country forestry reports, and social media, we compiled a global data set of 1683 case studies from over 125 NNT species, covering 44 countries, all continents but Antarctica, and seven biomes. Using different meta-analysis techniques, we found that, while NNTs increase most RES (e.g. climate regulation, soil erosion control, fertility and formation), they decrease PES (e.g. NNTs contribute less than native trees to global timber provision). Also, they have different effects on CES (e.g. increase aesthetic values but decrease scientific interest), and no effect on the EDS considered. NNT effects on each ecosystem (dis)service showed a strong context dependency, varying across NNT types, biomes and socio-economic conditions. For instance, some RES are increased more by NNTs able to fix atmospheric nitrogen, and when the ecosystem is located in low-latitude biomes; some CES are increased more by NNTs in less-wealthy countries or in countries with higher gross domestic products. The effects of NNTs on several ecosystem (dis)services exhibited some synergies (e.g. among soil fertility, soil formation and climate regulation or between aesthetic values and pollen allergenicity), but also trade-offs (e.g. between fire regulation and soil erosion control). Our analyses provide a quantitative understanding of the complex synergies, trade-offs and context dependencies involved for the effects of NNTs that is essential for attaining a sustained provision of ecosystem services.

Convergent xylem widening among organs across diverse woody seedlings.

Xylem conduit diameter (Dmax ) of woody angiosperm adults scales with plant size and widens from the stem apex downwards. We hypothesized that, notwithstanding relative growth rate (RGR), growth form or leaf habit, woody seedling conduit Dmax scales linearly with plant size across species; this scaling should be applicable to all vegetative organs, with consistent conduit widening from leaf via stem to main root and coupling with whole-leaf area and whole-stem xylem area. To test these hypotheses, organ-specific xylem anatomy traits and size-related traits in laboratory-grown seedlings were analyzed across 55 woody European species from cool-temperate and Mediterranean climates. As hypothesized, conduit Dmax of each organ showed similar scaling with plant size and consistent basipetal widening from the leaf midvein via the stem to the main root across species, independently of growth form, RGR and leaf habit. We also found a strong correlation between Dmax and average leaf area, and between stem xylem area and whole-plant leaf area. We conclude that seedlings of ecologically wide-ranging woody species converge in their allometric scaling of conduit diameters within and across plant organs. These relationships will contribute to modeling of water transport in woody vegetation that accounts for the whole life history from the trees' regeneration phase to adulthood.

Pollution Assessment of the Biobío River (Chile): Prioritization of Substances of Concern Under an Ecotoxicological Approach.

The water demand for human activities is rapidly increasing in developing countries. Under these circumstances, preserving aquatic ecosystems should be a priority which requires the development of quality criteria. In this study we perform a preliminary prioritization of the risky substances based on reported ecotoxicological studies and guidelines for the Biobío watershed (Central Chile). Our specific aims are (1) reviewing the scientific information on the aquatic pollution of this watershed, (2) determining the presence and concentration of potential toxic substances in water, sediment and effluents, (3) searching for quality criteria developed by other countries for the selected substances and (4) prioritizing the most risky substances by means of deterministic ecotoxicological risk assessment. We found that paper and mill industries were the main sources of point pollution, while forestry and agriculture were mostly responsible for non-point pollution. The most risky organic substances in the water column were pentachlorophenol and heptachlor, while the most relevant inorganic ones were aluminum, copper, unionized ammonia and mercury. The most risky organic and inorganic substances in the sediment were phenanthrene and mercury, respectively. Our review highlights that an important effort has been done to monitor pollution in the Biobío watershed. However there are emerging pollutants and banned compounds-especially in sediments-that require to be monitored. We suggest that site-specific water quality criteria and sediment quality criteria should be developed for the Biobío watershed, considering the toxicity of mixtures of chemicals to endemic species, and the high natural background level of aluminum in the Biobío.

Comparing the Sexual Reproductive Success of Two Exotic Trees Invading Spanish Riparian Forests vs. a Native Reference.

A widely accepted hypothesis in invasion ecology is that invasive species have higher survival through the early stages of establishment than do non-invasive species. In this study we explore the hypothesis that the sexual reproductive success of the invasive trees Ailanthus altissima (Mill.) Swingle and Robinia pseudoacacia L. is higher than that of the native Fraxinus angustifolia Vahl., all three species coexisting within the riparian forests of Central Spain. We compared different stages of the early life cycle, namely seed rain, seed infestation by insects, seed removal by local fauna, seed germination under optimal conditions and seedling abundance between the two invasive trees and the native, in order to assess their sexual reproductive success. The exotic species did not differ from the native reference (all three species displaying high seed rain and undergoing seed losses up to 50% due to seed removal by the local fauna). Even if the exotic R. pseudoacacia showed a high percentage of empty and insect-parasited seeds along with a low seedling emergence and the exotic A. altissima was the species with more viable seeds and of higher germinability, no differences were found regarding these variables when comparing them with the native F. angustifolia. Unsuitable conditions might have hampered either seedling emergence and survival, as seedling abundance in the field was lower than expected in all species -especially in R. pseudoacacia-. Our results rather suggest that the sexual reproductive success was not higher in the exotic trees than in the native reference, but studies focusing on long-term recruitment would help to shed light on this issue.

Can the life-history strategy explain the success of the exotic trees Ailanthus altissima and Robinia pseudoacacia in Iberian floodplain forests?

Ailanthus altissima and Robina pseudoacacia are two successful invasive species of floodplains in central Spain. We aim to explain their success as invaders in this habitat by exploring their phenological pattern, vegetative and sexual reproductive growth, and allometric relations, comparing them with those of the dominant native tree Populus alba. During a full annual cycle we follow the timing of vegetative growth, flowering, fruit set, leaf abscission and fruit dispersal. Growth was assessed by harvesting two-year old branches at the peaks of vegetative, flower and fruit production and expressing the mass of current-year leaves, stems, inflorescences and infrutescences per unit of previous-year stem mass. Secondary growth was assessed as the increment of trunk basal area per previous-year basal area. A. altissima and R. pseudoacacia showed reproductive traits (late flowering phenology, insect pollination, late and long fruit set period, larger seeds) different from P. alba and other native trees, which may help them to occupy an empty reproductive niche and benefit from a reduced competition for the resources required by reproductive growth. The larger seeds of the invaders may make them less dependent on gaps for seedling establishment. If so, these invaders may benefit from the reduced gap formation rate of flood-regulated rivers of the study region. The two invasive species showed higher gross production than the native, due to the higher size of pre-existing stems rather than to a faster relative growth rate. The latter was only higher in A. altissima for stems, and in R. pseudoacacia for reproductive organs. A. altissima and R. pseudoacacia showed the lowest and highest reproductive/vegetative mass ratio, respectively. Therefore, A. altissima may outcompete native P. alba trees thanks to a high potential to overtop coexisting plants whereas R. pseudoacacia may do so by means of a higher investment in sexual reproduction.

The relative importance for plant invasiveness of trait means, and their plasticity and integration in a multivariate framework.

Functional traits, their plasticity and their integration in a phenotype have profound impacts on plant performance. We developed structural equation models (SEMs) to evaluate their relative contribution to promote invasiveness in plants along resource gradients. We compared 20 invasive-native phylogenetically and ecologically related pairs. SEMs included one morphological (root-to-shoot ratio (R/S)) and one physiological (photosynthesis nitrogen-use efficiency (PNUE)) trait, their plasticities in response to nutrient and light variation, and phenotypic integration among 31 traits. Additionally, these components were related to two fitness estimators, biomass and survival. The relative contributions of traits, plasticity and integration were similar in invasive and native species. Trait means were more important than plasticity and integration for fitness. Invasive species showed higher fitness than natives because: they had lower R/S and higher PNUE values across gradients; their higher PNUE plasticity positively influenced biomass and thus survival; and they offset more the cases where plasticity and integration had a negative direct effect on fitness. Our results suggest that invasiveness is promoted by higher values in the fitness hierarchy--trait means are more important than trait plasticity, and plasticity is similar to integration--rather than by a specific combination of the three components of the functional strategy.

Developmental changes in mesophyll diffusion conductance and photosynthetic capacity under different light and water availabilities in Populus tremula: how structure constrains function.

Finite mesophyll diffusion conductance (g(m) ) significantly constrains net assimilation rate (A(n) ), but g(m) variations and variation sources in response to environmental stresses during leaf development are imperfectly known. The combined effects of light and water limitations on g(m) and diffusion limitations of photosynthesis were studied in saplings of Populus tremula L. An one-dimensional diffusion model was used to gain insight into the importance of key anatomical traits in determining g(m) . Leaf development was associated with increases in dry mass per unit area, thickness, density, exposed mesophyll (S(mes) /S) and chloroplast (S(c) /S) to leaf area ratio, internal air space (f(ias) ), cell wall thickness and chloroplast dimensions. Development of S(mes) /S and S(c) /S was delayed under low light. Reduction in light availability was associated with lower S(c) /S, but with larger f(ias) and chloroplast thickness. Water stress reduced S(c) /S and increased cell wall thickness under high light. In all treatments, g(m) and A(n) increased and CO(2) drawdown because of g(m) , C(i) -C(c) , decreased with increasing leaf age. Low light and drought resulted in reduced g(m) and A(n) and increased C(i) -C(c) . These results emphasize the importance of g(m) and its components in determining A(n) variations during leaf development and in response to stress.

Establishment success of coexisting native and exotic trees under an experimental gradient of irradiance and soil moisture.

The exotic trees Ailanthus altissima, Robinia pseudoacacia, Acer negundo and Elaeagnus angustifolia coexist with the native trees Fraxinus angustifolia and Ulmus minor in river banks of central Spain. Similarly, the exotic trees Acacia dealbata and Eucalyptus globulus co-occur with the natives Quercus pyrenaica and Pinus pinaster in Northwest Spain. We aimed to identify the environmental conditions that favour or hamper the establishment success of these species. In spring 2008, seeds of the studied species were sown under an experimental gradient of light (100, 65, 35, 7% of full sunlight) combined with three levels of soil moisture (mean soil water potential = -0.97, -1.52 and -1.77 MPa.). During the first growing season we monitored seed emergence and seedling survival. We found that the effect of light on the establishment success was stronger than the effect of soil moisture. Both exotic and native species of central Spain showed a good performance under high light, A. negundo being the most shade tolerant. Water shortage diminished E. angustifolia and A. altissima success. Among NW Spain species, A. dealbata and P. pinaster were found to be potential competitors for colonizing high-irradiance scenarios, while Q. pyrenaica and E. globulus were more successful under moderate shade. High soil moisture favoured E. globulus but not A. dealbata establishment. These results contribute to understand some of the factors controlling for spatial segregation between coexisting native and exotic tree species, and can help to take decisions orientated to the control and management of these exotic species.

Leaf litter traits of invasive species slow down decomposition compared to Spanish natives: a broad phylogenetic comparison.

Leaf traits related to the performance of invasive alien species can influence nutrient cycling through litter decomposition. However, there is no consensus yet about whether there are consistent differences in functional leaf traits between invasive and native species that also manifest themselves through their "after life" effects on litter decomposition. When addressing this question it is important to avoid confounding effects of other plant traits related to early phylogenetic divergences and to understand the mechanism underlying the observed results to predict which invasive species will exert larger effects on nutrient cycling. We compared initial leaf litter traits, and their effect on decomposability as tested in standardized incubations, in 19 invasive-native pairs of co-familial species from Spain. They included 12 woody and seven herbaceous alien species representative of the Spanish invasive flora. The predictive power of leaf litter decomposition rates followed the order: growth form > family > status (invasive vs. native) > leaf type. Within species pairs litter decomposition tended to be slower and more dependent on N and P in invaders than in natives. This difference was likely driven by the higher lignin content of invader leaves. Although our study has the limitation of not representing the natural conditions from each invaded community, it suggests a potential slowing down of the nutrient cycle at ecosystem scale upon invasion.

Seasonal carbon storage and growth in Mediterranean tree seedlings under different water conditions.

In all Mediterranean-type ecosystems, evergreen and deciduous trees differing in wood anatomy, growth pattern and leaf habit coexist, suggesting distinct adaptative responses to environmental constraints. This study examined the effects of summer water stress on carbon (C) storage and growth in seedlings of three coexisting Mediterranean trees that differed in phenology and wood anatomy characteristics: Quercus ilex subsp. ballota (Desf.) Samp., Quercus faginea Lam. and Pinus halepensis L. Seedlings were subjected to two levels of watering during two consecutive summers and achieved a minimum of -0.5 and -2.5 MPa of predawn water potential in the control and water stress treatment, respectively. Both Quercus species concentrated their growth in the early growing season, demanding higher C in early spring but replenishing C-stores in autumn. These species allocated more biomass to roots, having larger belowground starch and lipid reserves. Quercus species differed in seasonal storage dynamics from P. halepensis. This species allocated most of its C to aboveground growth, which occurred gradually during the growing season, leading to fewer C-reserves. Soluble sugar and starch concentrations sharply declined in August in P. halepensis, probably because reserves support respiration demands as this species closed stomata earlier under water stress. Drought reduced growth of the three species, mainly in Q. faginea and P. halepensis, but not C-reserves, suggesting that growth under water stress conditions is not limited by C-availability.

Flowering phenology of invasive alien plant species compared with native species in three Mediterranean-type ecosystems.

BACKGROUND AND AIMS: Flowering phenology is a potentially important component of success of alien species, since elevated fecundity may enhance invasiveness. The flowering patterns of invasive alien plant species and related natives were studied in three regions with Mediterranean-type climate: California, Spain and South Africa's Cape region. METHODS: A total of 227 invasive-native pairs were compared for seven character types across the regions, with each pair selected on the basis that they shared the same habitat type within a region, had a common growth form and pollination type, and belonged to the same family or genus. KEY RESULTS: Invasive alien plant species have different patterns of flowering phenology from native species in the three regions. Whether the alien species flower earlier, later or at the same time as natives depends on the climatic regime in the native range of the aliens and the proportion of species in the invasive floras originating from different regions. Species invading at least two of the regions displayed the same flowering pattern, showing that flowering phenology is a conservative trait. Invasive species with native ranges in temperate climates flower earlier than natives, those from Mediterranean-type climates at the same time, and species from tropical climates flower later. In California, where the proportion of invaders from the Mediterranean Basin is high, the flowering pattern did not differ between invasive and native species, whereas in Spain the high proportion of tropical species results in a later flowering than natives, and in the Cape region early flowering than natives was the result of a high proportion of temperate invaders. CONCLUSIONS: Observed patterns are due to the human-induced sympatry of species with different evolutionary histories whose flowering phenology evolved under different climatic regimes. The severity of the main abiotic filters imposed by the invaded regions (e.g. summer drought) has not been strong enough (yet) to shift the flowering pattern of invasive species to correspond with that of native relatives. It does, however, determine the length of the flowering season and the type of habitat invaded by summer-flowering aliens. Results suggest different implications for impacts at evolutionary time scales among the three regions.

Water relations of seedlings of three Quercus species: variations across and within species grown in contrasting light and water regimes.

We compared seedling water relations of three Mediterranean Quercus species (the evergreen shrub Q. coccifera L., the evergreen tree Q. ilex L. subsp. ballota (Desf.) Samp. and the deciduous or marcescent tree Q. faginea L.). We also explored seedling potential for acclimation to contrasting growing conditions. In March, 1-year-old seedlings of the three species were planted in pots and grown outdoors in a factorial combination of two irrigation regimes (daily (HW) and alternate day watering (LW)) and two irradiances (43 and 100% of full sunlight). At the end of July, predawn and midday water potentials (Psi(pd), Psi(md)) were measured, and pressure-volume (P-V) curves were obtained for mature current-year shoots. Species exhibited similar Psi(pd) and Psi(md) values, but differed in leaf morphology and water relations. The evergreens possessed larger leaf mass per area (LMA) and were able to maintain positive turgor pressure at lower water potentials than the deciduous species because of their lower osmotic potential at full turgor. However, the three species had similar relative water contents at the turgor loss point because Q. faginea compensated for its higher osmotic potential with greater cell wall elasticity. Values of Psi(pd) had a mean of -1.12 MPa in LW and -0.63 MPa in HW, and Psi(md) had a mean of -1.13 MPa in full sunlight and -1.64 MPa in shade, where seedlings exhibited lower LMA. However, the P-V curve traits were unaffected by the treatments. Our results suggest that Q. faginea seedlings combine the water-use characteristics of mesic deciduous oak and the drought-tolerance of xeric evergreen oak. The ability of Q. coccifera to colonize drier sites than Q. ilex was not a result of higher drought tolerance, but rather may be associated with other dehydration postponement mechanisms including drought-induced leaf shedding. The lack of treatment effects may reflect a relatively low contrast between treatment regimes, or a low inherent responsiveness of these traits in the study species, or both.

Interactive effects of shade and irrigation on the performance of seedlings of three Mediterranean Quercus species.

Shade and irrigation are frequently used to increase the success of Mediterranean Quercus spp. plantations. However, there is controversy about the combined effects of these treatments on plant performance. We assessed the effects of two irradiances (full sunlight and moderate shade) and two summer watering regimes (high (daily) and low (alternate days)) on leaf and whole-plant traits of 1-year-old seedlings of Quercus coccifera, Q. ilex subsp. ballota and Q. faginea grown outdoors for 8.5 months. Leaf traits included measures of morphology, nitrogen concentration, gas exchange and photochemical efficiency, and measures of whole-plant traits included biomass allocation patterns, growth phenology, across-summer leaf area change and relative growth rate (RGR). Moderate shade reduced leaf mass per area, increased photochemical efficiency, maximum carbon assimilation rate (Amax) and allocation to leaves, and prolonged the growing period in one or more of the species. Daily watering in summer increased Amax of Q. ilex and prolonged the growing period of Q. ilex and Q. faginea. Both treatments tended to increase RGR. The effect of shade was greater in the low-watering regime than in the high-watering regime for two of the 15 studied traits, with treatment effects being independent for the remaining 13 traits. Leaf nitrogen and the ability to maintain leaf area after the arid period, rather than biomass allocation traits, explained the variation in seedling RGR. Trait responsiveness to the treatments was low and similar among species and between study scales, being unexpectedly low in Q. faginea leaves. This may be because selective pressures on leaf plasticity act differently in deciduous and evergreen species. We conclude that moderate shade and daily summer watering enhance the performance of Mediterranean Quercus seedlings through species-specific mechanisms.