Differential effect of drought regimes on the seedling performance of six floodplain grassland species

• The performance of seedlings is crucial for the survival and persistence of plant populations. Although drought frequently occurs in floodplains and can cause seedling mortality, studies on the effects of drought on seedlings of floodplain grasslands are scarce. We tested the hypotheses that drought reduces aboveground biomass, total biomass, plant height, number of leaves, leaf area and specific leaf area (SLA), and increases root biomass and root‐mass fraction (RMF) and that seedlings from species of wet floodplain grasslands are more affected by drought than species of dry grasslands.
• In a greenhouse study, we exposed seedlings of three confamilial pairs of species (Pimpinella saxifraga, Selinum carvifolia, Veronica teucrium, Veronica maritima, Sanguisorba minor, Sanguisorba officinalis) to increasing drought treatments. Within each plant family, one species is characteristic of wet and one of dry floodplain grasslands, confamilial in order to avoid phylogenetic bias of the results.
• In accordance with our hypotheses, drought conditions reduced aboveground biomass, total biomass, plant height, number of leaves and leaf area. Contrary to our hypotheses, drought conditions increased SLA and decreased root biomass and RMF of seedlings. Beyond the effects of the families, the results were species‐specific (V. maritima being the most sensitive species) and habitat‐specific. Species indicative of wet floodplain grasslands appear to be more sensitive to drought than species indicative of dry grasslands.
• Because of species‐ and habitat‐specific responses to reduced water availability, future drought periods due to climate change may severely affect some species from dry and wet habitats, while others may be unaffected.

     

Invasion Success in a Marginal Habitat: An Experimental Test of Competitive Ability and Drought Tolerance in Chromolaena odorata

Climatic niche models based on native-range climatic data accurately predict invasive-range distributions in the majority of species. However, these models often do not account for ecological and evolutionary processes, which limit the ability to predict future range expansion. This might be particularly problematic in the case of invaders that occupy environments that would be considered marginal relative to the climatic niche in the native range of the species. Here, we assess the potential for future range expansion in the shrub Chromolaena odorata that is currently invading mesic savannas (>650 mm MAP) in South Africa that are colder and drier than most habitats in its native range. In a greenhouse experiment we tested whether its current distribution in South Africa can be explained by increased competitive ability and/or differentiation in drought tolerance relative to the native population. We compared aboveground biomass, biomass allocation, water use efficiency and relative yields of native and invasive C. odorata and the resident grass Panicum maximum in wet and dry conditions. Surprisingly, we found little differentiation between ranges. Invasive C. odorata showed no increased competitive ability or superior drought tolerance compared to native C. odorata. Moreover we found that P. maximum was a better competitor than either native or invasive C. odorata. These results imply that C. odorata is unlikely to expand its future range towards more extreme, drier, habitats beyond the limits of its current climatic niche and that the species’ invasiveness most likely depends on superior light interception when temporarily released from competition by disturbance. Our study highlights the fact that species can successfully invade habitats that are at the extreme end of their ranges and thereby contributes towards a better understanding of range expansion during species invasions.     

Root growth of different oak provenances in two soils under drought stress and air warming conditions

Background and aims

Oaks are considered to be drought- and thermo-tolerant tree species. Nevertheless, species and provenances may differ in their ecological requirements. We hypothesised that (i) provenances from xeric sites are better adapted to drought than those from more humid sites, (ii) oaks direct root growth towards resource-rich layers, and (iii) air-warming promotes root growth.

Methods

To test different provenances of Quercus robur, Q. petraea and Q. pubescens, we conducted a model ecosystem experiment with young trees, grown on acidic and calcareous soil, subjected to drought, air warming, the combination of warming and drought, and a control.

Results

The results were only in partial agreement with the first hypothesis. As expected the provenances originating from drier sites produced more biomass than those from more humid sites under drought conditions. Surprisingly, however, they reacted more sensitive to water limiting conditions, as they produced also substantially more biomass under well-watered conditions. The drought treatment reduced root mass substantially in the upper soil. In agreement with the second hypothesis this led to a shift in the centre of root mass to lower depth, where water was still more available than closer to the soil surface. In contrast to the third hypothesis, the air-warming treatment, which was very mild however compared to climate change scenarios, had no significant effects on root growth.

Conclusions

Given that the provenances from drier sites showed more biomass loss at water limiting conditions than provenances from more humid sites, it remains questionable whether provenances from drier sites are better suited for a future climate.     

Breaking dormancy during flood and drought: sublethal growth and physiological responses of three emergent wetland herbs used in bioretention basins

Bioretention basins are man-made topographic depressions designed to collect and retain surface water runoff. In most cases these basins are used to prevent flooding and/or remove environmentally harmful pollutants and sediments from entering natural aquatic systems. To maximize environmental benefits, these systems are often planted with flood-tolerant wetland hydrophytes that are capable of withstanding extended periods of drought. Unfortunately, little is known about how these plants respond to extreme hydrologies while breaking seasonal dormancy. The purpose of this study was to better understand the morphological and physiological responses of three wetland species (Pontederia cordata L., Saururus cernuus L., and Schoenoplectus tabernaemontani C.C. Gmel. Palla) often used in bioretention basins while emerging from dormancy in either flooded or drought conditions. Results indicate that only S. tabernaemontani was affected morphologically by drought with lower leaf area and aboveground biomass. While significant reductions in stomatal surface indices were also observed in drought-treated S. tabernaemontani, all three species had reductions in stomatal conductance (g) when grown in drier soils. Moreover, drought conditions promoted decreases in leaf water potential (Ψ _leaf) for all three species, and reductions in tissue water content (θ) for P. cordata and S. tabernaemontani. Based on the overall morphological and physiological responses, S. cernuus maintained the lowest productivity, and appeared to be the best suited for tolerating sustained soil water deficits. If high plant productivity is desired, however, S. tabernaemontani was able to maintain high plant growth while making necessary modifications that facilitated greater drought-resistance.     

Sporadic rainy events are more critical than increasing of drought intensity for woody species recruitment in a Mediterranean community

The understanding of the impact of extreme climatic events under a global climate change scenario is crucial for the accurate forecast of future plant community dynamics. We have experimentally assessed the effect of drier and wetter summer conditions on the recruitment probabilities and the growth of seedlings from eight woody species representative of the most important functional groups in the community, pioneer shrubs, mid-successional shrubs and trees, across the main habitats in the study area (open habitat, shrubland, and forest). Our hypothesis proposes that wet summer conditions would represent a good opportunity for tree species regeneration, enhancing both forest maintenance and expansion. A drier summer scenario, on the other hand, would limit forest regeneration, and probably hinder the colonization of nearby habitats. We found a habitat effect on the emergence, survival, and final biomass, whereas different climate scenarios affected seedling survival and biomass. A wet summer boosted growth and survival, whereas greater drought reduced survival only in some cases. These results were modulated by the habitat type. Overall, shrub species presented higher survival and growth and were less affected by more severe drought, whereas some tree species proved to be extremely dependent on wet summer conditions. We conclude that the reduction in frequency of wet summers predicted for the coming decades in Mediterranean areas will have greater consequences for species recruitment than will increased drought. The different response of the species from the various functional groups has the potential to alter the composition and dominance of future plant communities.     

Waterlogging Stress and Ethylene Production in the Dune Slack Plant, Scirpus americanus

The sedge, Scirpus americanus Pers., grows in dune slacks andother freshwater and brackish water wetland communities. Whenwaterlogged in a greenhouse, the concentration of ethylene increased4-fold in stems of S. americanus plants. This increase was associatedwith a decrease in plant height and an increase in aerenchymaas exhibited under waterlogged conditions. Endogenous ethyleneproduction in S. americanus was compared to that in anotherdune slack species, Panicum amarulum, and also to Spartina aherniflorafrom a salt marsh. These species did not respond by increasingendogenous ethylene upon waterlogging. In the field, a 16 cmrainfall significantly increased the endogenous ethylene productionin S. americanus. As the water table subsided the concentrationof accumulated ethylene in stem tissue decreased. Exposure ofS. americanus to exogenous ethylene inhibited stem extensionand increased aerenchyma formation, thus linking ethylene tothe morphological characteristics of waterlogged plants of thisspecies. These experiments support the hypothesis that ethylenemodulates S. americanus morphology in natural waterlogged environmentsand may be of importance in adapting this species to life inthe wetland environment. Key words: Dune slack, waterlogging, ethylene     

Temporal variations and spatial patterns in saline and waterlogged peat fields: 1. Survival and growth of salt marsh graminoids

A cutover bog contaminated with seawater in New Brunswick, Canada remained barren 5 years after peat extraction operations ceased despite the proximity of natural seed sources. The aim of the study was to identify abiotic stresses impeding plant establishment and test transplanting of salt-tolerant wetland plants. The site consisted of long cambered rectangular fields that sloped down (2%) to the drainage ditches on both sides. Across this slope, zones were delineated based on moisture gradient as: Up-areas (drier), Mid-areas (moist) and Low-areas (wet). Juncus balticus was transplanted to these zones in August 2004 whereas Spartina pectinata was reintroduced in June 2005. Plant material was collected from nearby marshes. Survival of J. balticus in August 2005 was poorest at the Low-areas probably because of the early season flooded conditions of that zone. S. pectinata survival in June 2006 was good in all zones having better adaptation to early season waterlogged conditions. Early season waterlogged conditions resulted from a perched water table (May–June) and were alleviated only upon the complete thaw of the frozen peat layer on 8 July. Thereafter, important changes in peat characteristics occurred: lowered water table depths that increased redox potentials, decreased moisture content (θ) that increased dry bulk density (ρ), and increased electrical conductivity (EC) that decreased pH. Waterlogged conditions were of greater magnitude and duration at the Low-areas but salinity levels were highest in the Up-areas and Mid-areas. We conclude that local conditions of waterlogging (especially in the Low-areas) and high salinity and low pH (notably in the Up- and Mid-areas) favoured the survival of S. pectinata in all areas and J. balticus in Up- and Mid-areas only.     

High competitiveness of a resource demanding invasive acacia under low resource supply

Mechanisms controlling the successful invasion of resource demanding species into low-resource environments are still poorly understood. Well-adapted native species are often considered superior competitors under stressful conditions. Here we investigate the competitive ability of the resource demanding alien Acacia longifolia, which invades nutrient-poor Mediterranean sand dunes such as in coastal areas of Portugal. We explore the hypothesis that drought may limit invasion in a factorial competition experiment of the alien invasive versus two native species of different functional groups (Halimium halimifolium, Pinus pinea), under well-watered and drought conditions. Changes in biomass, allocation pattern, and N-uptake-efficiency (via ¹⁵N-labeling) indicated a marked drought sensitivity of the invader. However, highly efficient drought adaptations of the native species did not provide a competitive advantage under water limiting conditions. The competitive strength of H. halimifolium towards the alien invader under well-watered conditions turned into a positive interaction between both species under drought. Further, low resource utilization by native species benefited A. longifolia by permitting continued high nitrogen uptake under drought. Hence, the N-fixing invader expresses low plasticity by continuous high resource utilization, even under low resource conditions. The introduction of novel traits into a community like N-fixation and high resource use may promote A. longifolia invasiveness through changes in the physical environment, i.e., the water and nutrient cycle of the invaded sand dune system, thereby potentially disrupting the co-evolved interactions within the native plant community.     

Propagule pressure and environmental conditions interact to determine establishment success of an invasive plant species,glossy buckthorn (<Emphasis Type="Italic">Frangula alnus</Emphasis>), across five different wetland habitat types

Many invasive plant species are able to establish within a wide range of community types. This establishment success depends on high propagule pressure and successful recruitment of seedlings following propagule dispersal into receptive environments. This study aimed to investigate interactions between propagule pressure and environmental resistance to seedling recruitment of the invasive shrub, glossy buckthorn (Frangula alnus Mill.), over a range of wetland habitat types. We measured propagule deposition using seed traps and recruitment success using sown plots, while characterizing vegetation and abiotic environmental conditions in five adjacent wetland habitat types. Drier habitats, which included Cedar Swamp, Shrub Carr, and String, had lower resistance to buckthorn establishment than the wetter Flark and Cattail Marsh. The drier habitats supported more woody species and provided more raised hummock surfaces essential for successful buckthorn recruitment and establishment. Propagule pressure was also higher in dry habitats that supported higher densities of adult glossy buckthorn, while long-distance dispersal into areas with low adult density was uncommon. Natural recruitment was highest in sites with intense propagule pressure, but experimental sowing of seeds demonstrated that buckthorn establishes in wet sites with higher resistance if propagule pressure is increased and seeds are deposited on hummocks. This study demonstrates the affinity of glossy buckthorn for drier wetland sites, and provides empirical evidence that environmental resistance can be overcome by higher propagule pressure.     

Differential impact of hotter drought on seedling performance of five ecologically distinct pine species

Increasing temperature and drought intensity is inducing the phenomenon of the so-called “hotter drought”, which is expected to increase in frequency over the coming decades across many areas of the globe, and is expected to have major implications for forest systems. Consequences of hotter drought could be especially relevant for closely related species overlapping their distributions, since differences in response can translate into range shifts. We assessed the effect of future climatic conditions on the performance of five ecologically distinct pine species common in Europe: Pinus halepensis, P. pinaster, P. nigra, P. sylvestris and P. uncinata. We hypothesised that Mediterranean species inhabiting dry, low-elevation sites will be less affected by the expected warming and drought increase than species inhabiting cold-wet sites. We performed a controlled conditions experiment simulating current and projected temperature and precipitation, and analysed seedling responses in terms of survival, growth, biomass allocation, maximum photochemical efficiency (F _v/F _m) and plant water potential (Ψ). Either an increase in temperature or a reduction in water input alone reduced seedling performance, but the highest impact occurred when these two factors acted in combination. Warming and water limitation reduced Ψ, whereas warming alone reduced biomass allocation to roots and F _v/F _m. However, species responded differentially to warmer and drier conditions, with lowland Mediterranean pines (P. halepensis and P. pinaster) showing higher survival and performance than mountain species. Interspecific differences in response to warmer, drier conditions could contribute to changes in the relative dominance of these pine species in Mediterranean regions where they co-occur and a hotter, drier climate is anticipated.     

Occurrence patterns of facilitation by shade along a water gradient are mediated by species traits

In disturbed habitats, shade often has facilitative effects on plants by ameliorating water and thermal stresses. Facilitation by shade tends to increase as water availability decreases. At the same time, several studies have suggested that facilitation by shade is not affected by water status or collapses under extremely dry conditions. We hypothesized that traits of beneficiary plants, specifically, the flexibility in the allocation of biomass between shoots and roots, would mediate variation in the relationship between facilitation by shade and water status. To test this hypothesis, we examined the responses of two bog species to shade under various water conditions in a post-mined peatland. The seeds of Rhynchospora alba and Moliniopsis japonica were sown under three water levels (dry: 53% peat water content, wet: 77%, and control: 71%) × two shading levels (50% shaded and unshaded). The survival, biomass, and biomass allocation between the shoots and roots of the two species were monitored for two years. Shade increased the survival and biomass of both species. However, the facilitation of R. alba by shade was independent of water level, whereas the strength of the facilitative effects on M. japonica increased as water content decreased. R. alba preferentially allocated biomass to roots under dry conditions and was highly drought tolerant. M. japonica did not alter the allocation of its biomass in response to either shade or water level and was drought intolerant. Our results suggest that flexibility in biomass allocation of beneficiary plants mediates occurrence patterns of facilitation by shade along a water gradient. The facilitation of species with inflexible biomass allocation by shade through the amelioration of water stress increases as water availability decreases, whereas the facilitation of species with flexible biomass allocation is independent of water status. Such species-specific facilitation would promote the coexistence of diverse species in a community.     

Effects of climate‐induced increases in summer drought on riparian plant species: a meta‐analysis

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Effect of an Epichloë endophyte on adaptability to water stress in Festuca sinensis

As water stress, including drought and waterlogging, can severely affect plant growth, this study investigated the effects of an endophyte from the genus Epichloë on two different ecotypes of Festuca sinensis grass under five soil water conditions in a controlled greenhouse experiment. Changes in F. sinensis plants grown with (E+) and without the endophyte (E−) were evaluated as they were subjected to different water treatments (20%, 35%, 50%, 65% and 80% relative saturation moisture content, RSMC). Growth parameters such as plant height, number of tillers, blade width, stem diameter, root length, total biomass, root-shoot ratio and relative water content were determined. The results showed that drought and waterlogging significantly (P < 0.05) inhibited the growth of F. sinensis. The presence of the endophyte significantly (P < 0.05) increased plant growth and root-shoot ratio under drought and waterlogged conditions. In addition, the plant height, number of tillers, blade width, stem diameter and total biomass in seedlings of both ecotypes reached the maximum at 65% RSMC, which suggests the optimal water condition. These findings also show that moderate drought (35% and 50% RSMC) could promote root growth of grass seedlings. Therefore, endophytic infections can result in enhanced host plant resistance to drought and waterlogged conditions.     

Evaluating the realized niche and plant–water relations of wetland species using experimental transplants

Wet meadows are a common focus of wetland restoration efforts, and the species within them often exist within a restricted range of water levels. Unfortunately, many restored wetlands have higher water levels and more open water than naturally occurring reference wetlands, and many are invaded and dominated by species of Typha. Most studies evaluating the optimal water level for plant species use observational methods, yet experimental methods are required to understand the breadth of a species’ niche. We used experimental transplants of Carex pellita, a common wet meadow sedge used in restoration in the interior of the USA, and Typha latifolia, a species of cattail which invades many restored wetlands, to test whether higher water levels in a restored wetland were prohibitive to the target sedge species. Physiological and growth measurements were collected on both species. We found that C. pellita grew as well or better when transplanted into the ponded water levels, while T. latifolia had reduced growth when transplanted into the relatively drier meadow conditions. Interestingly, C. pellita was able to adjust its Turgor Loss Point in response to changing water levels. Only recently the assumption of a constant Turgor Loss Point for each species has been questioned. Our results provide evidence that wet meadow species have a broader hydrologic niche than previously thought, and their ability to make physiological adjustments in response to changing water levels may allow them to thrive in areas with widely varying water levels.

     

Facilitation vs. competition: Does interspecific interaction affect drought responses in Sphagnum?

The stress-gradient hypothesis (SGH) predicts that the relative importance of competition decreases and facilitation increases with an increase in abiotic stress. In peatlands, Sphagnum faces the threat of drought and differentiates into hummock species (drought-tolerant) and hollow species. Whether interspecific interaction affects the influence of drought on bryophyte composition in peatlands is unknown. We established an experiment by simulating drought and building bryophyte communities with two hummock species (S. palustre and S. capillifolium) and one hollow species (S. fallax). In all three species, drought decreased biomass production, height increment and side-shoot production. Sphagnum stores water in the hyaline cells, and leaf hyaline cell percentage (HCP) in the two hummock species increased with drought while no effect was found in S. fallax, suggesting that adjusting HCP is not an effective response to drought for the hollow species. Morphological traits and carbon and nitrogen contents in hummock species responded more to drought than in the hollow species, indicating a rapid response in phenotypic plasticity is an important strategy to resist drought in the hummock species. The presence of neighboring Sphagnum species, rather than drought, decreased carbon content for all three species. All three bryophytes showed interaction between drought and neighbor in two or more plant traits. Our study, however, did not support SGH, and there were no changes from competition under wet to facilitation under dry treatments in any of the six species combinations. On the contrary, when S. fallax was the target species, a change from facilitation under wet to competition under dry treatments was observed. The results suggest that hummock species can facilitate hollow species in wet environments but they could suppress hollow species under drought conditions by competing for water resources. Both drought and strong competition are the probable reasons why hollow species rarely grow in hummocks.     

Tree growth,mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought

A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests.     

Evidence for plant traits driving specific drought resistance. A community field experiment

Drought is known to be a major bottleneck for woody-community recruitment. The species-specific resistance to drought and factors involved in its variation are of special interest to forecast community fate. We performed an experiment under natural field conditions from winter 2006 to end summer 2008 to investigate the structural responses of woody saplings to nine combinations of light (three habitats differing in plant cover: forest, shrubland, and open) and water (three climate scenarios: drier, current and wetter summers). Our working hypothesis is that plant strategies to cope with drought are determined by habitat characteristics and/or variation of plant traits, and that these different strategies may determine community composition and dynamics. Eight woody species with different life forms and successional stages (trees, mid-successional shrubs, and pioneer shrubs) were selected for the comparison, including therefore species representative of the entire woody community. We explored drought resistance at the population level (D_S), the relations of the different morphological traits to D_S, and the potential importance for plants of inter-specific trade-offs. D_S ranged from 0% to 99% for the different species, depending on the habitat. Some structural traits were found to be related to D_S: positively with total biomass and leaf mass ratio (LMR) and negatively with leaf area-root mass ratio (LARMR). Contrary to previous studies, the present work revealed no evidence of trade-offs, such as survival in forest vs. growth in open, or growth in forest vs. growth in open. Accordingly, some species with low D_S values (Acer opalus and Pinus sylvestris) would be threatened under the future climate conditions, while species having structural characteristics to increase their resistance under expected dry years in coming decades (i.e., high LMR and total biomass or low LARMR, such as Quercus ilex and the shrub Cytisus scoparius) might enhance their recruitment probabilities. Thus, species-specific plant traits, and their effect on D_S, may filter future community assemblages.     

<Emphasis Type="Italic">Pinus nigra</Emphasis> Arn. ssp <Emphasis Type="Italic">salzmannii</Emphasis> early recruitment and initial seedling growth in warmer and drier locations: the role of seed and soil provenance

Current decreasing precipitation and increasing temperatures promote the likelihood of extreme drought events and may alter the recruitment capacities of tree species. Spanish black pine (Pinus nigra ssp. salzmannii) initial recruitment is being one of the most affected pine species by changing conditions with alterations in the future species distribution. In this context, a cross-exchange experiment was implemented using an outdoor nursery located in a warmer and drier location for testing different Spanish Black pine seeds and soil provenance combinations in relation to early recruitment and initial seedling growth. Soil and seeds were collected at a high (HA, 1641 m.a.s.l.) and low (LA, 1099 m.a.s.l.) altitude in Cuenca Mountains (Spain). Then, a cross-sown experiment using HA and LA soils and seeds was set up in an outdoor nursery, which is located in Albacete (704 m.a.s.l.). Soil quality, seedling emergence, seedling survival, initial seedling growth and total seedling dry mass were measured after one year. We found higher seed emergence and seedling survival by combining LA soil with LA seeds or HA soil with HA seeds. Seedlings from LA seeds with both soil origins and seedlings from HA seeds with LA soils allocated more biomass to roots than seedlings from HA sites growing in HA soils under drier and warmer conditions. These results support the idea that autochthonous provenances have the potential to adapt to changing climatic conditions in their habitats.     

Effect of wet and dry cycles in calcareous soil on mineral nutrient uptake of two grasses,Agrostis stolonifera L. and Festuca ovina L.

Shoot uptake of mineral nutrients (Ca, Cu, Fe, K, mg, Mn, P, S, Zn) by Agrostis stolonifera L. was compared with Festuca ovina L. under wet and dry cycles. Such conditions are typical for A. stolonifera sites, whereas F. ovina is growing mostly on consistently drier and better-drained soils. Plants were grown in a glasshouse, at controlled temperature and light conditions, using two moisture regimes, one constant at 60% WHC (water holding capacity), one wet/dry fluctuating between 35 and 100% WHC. Above ground and total biomass production was lower under wet/dry treatment than at constant water regime in F. ovina, but did not differ between regimes in A. stolonifera. Shoot uptake of most elements was severely reduced in F. ovina at the wet/dry regime. Shoot uptake and concentrations of most elements studied (Cu, K, Mn, P, S, Zn) were lower (p<0.05) under wet/dry treatment than at constant regime in A. stolonifera and tended to be lower also of Fe and Mg. Differences in biomass production observed are consistent with field evidence that A. stolonifera grows in sites which are periodically flooded but may become quite dry during other periods, and that F. ovina is limited to sites which are consistently drier and better drained. Evidence from the present study, however, does not support any view that alternating wet and dry cycles, as typical of A. stolonifera field sites, would be beneficial to nutrient acquisition of this species but that biomass production may develop normally at the lower uptake of most mineral nutrients measured under the wet/dry regime. Such regimes are decidedly unfavourable to both growth and nutrient acquisition of F. ovina.     

Developmental instability as an index of adaptation to drought stress in a Mediterranean oak

An increase in temperature and water deficits caused by the ongoing climate change might lead to a decline growth rates and threaten the persistence of tree species in drought-prone areas within the Mediterranean Basin. Developmental instability (the error in development caused by stress) may provide an index of the adaptability of woody plants to withstand climatic stressors such as water shortage. This study evaluated the effects of drought stress on growth variables in three stands of a Mediterranean oak (Quercus faginea) exposed to differing climatic conditions (xeric, mesic and cooler) along an altitudinal gradient in northeastern Spain, in two climatically contrasting years (wet and dry years). Two indices of developmental instability, fluctuating and translational asymmetries, which reflect environmental stress, were measured in leaves and current-year shoots, respectively. We also measured branch biomass and fractal complexity of branches as indicators of the species’ performance. After a period of drought the individuals’ at the most xeric site presented lower developmental instability and less branch biomass than did the individuals from the mesic and cooler sites. We interpret that difference as an adaptive response to drought which reflects a trade-off between maintenance of homeostasis and growth when water is scarce. The study demonstrated that developmental instability constitutes a useful index to assess the degree of adaptation to stressful environmental conditions. The assessment of developmental instability in sites and years with contrasting climatic conditions provides a means of quantifying the capacity of plants to develop plastic adaptive responses to climatic stress.