Semi-arid Mediterranean Ecosystems Research Articles

Soil microbial respiration beneath Stipa tenacissima L. and in surrounding bare soil

Open steppes dominated by Stipa tenacissima L. constitute one of the most representative ecosystems of the semi-arid zones of Eastern Mediterranean Basin (Iberian Peninsula, North of Africa). Ecosystem functioning of these steppes is strongly related to the spatial pattern of grass tussocks. Soils beneath Stipa tenacissima L. grass show different fertility and different microclimatic conditions than in surrounding bare soil. The objective of this study was to assess the effect of Stipa tenacissima L. on the key soil microbial activities under controlled incubation conditions (basal and potential respiration). Basal and potential microbial respirations in the soils beneath Stipa tenacissima L. were, in general, not significantly different from the bare soils. The differences were less than 10%. Significantly less ethylene produced by microbial activity in soils beneath Stipa tenacissima L. after the addition of glucose could indicate the dependence of rhizospheric microbial communities on available carbon compounds. It can be concluded, that the soil respiration in semi-arid Mediterranean ecosystems is not necessarily associated with the patchy plant distribution and that some microbial activities characteristics can be unexpectedly homogenous.

Ecological functioning in grass-shrub Mediterranean ecosystems measured by eddy covariance.

Climate change may alter ecosystem functioning, as assessed via the net carbon (C) exchange (NEE) with the atmosphere, composed of the biological processes photosynthesis (GPP) and respiration (R(eco)). In addition, in semi-arid Mediterranean ecosystems, a significant fraction of respired CO2 is stored in the vadose zone and emitted afterwards by subsoil ventilation (VE), contributing also to NEE. Such conditions complicate the prediction of NEE for future change scenarios. To evaluate the possible effects of climate change on annual NEE and its underlying processes (GPP, R(eco) and VE) we present, over a climate/altitude range, the annual and interannual variability of NEE, GPP, R(eco) and VE in three Mediterranean sites. We found that annual NEE varied from a net source of around 130 gC m(-2) in hot and arid lowlands to a net sink of similar magnitude for alpine meadows (above 2,000 m a.s.l) that are less water stressed. Annual net C fixation increased because of increased GPP during intermittent and several growth periods occurring even during winter, as well as due to decreased VE. In terms of interannual variability, the studied subalpine site behaved as a neutral C sink (from emission of 49 to fixation of 30 gC m(-2) year(-1)), with precipitation as the main factor controlling annual GPP and R(eco). Finally, the importance of VE as 0-23% of annual NEE is highlighted, indicating that this process could shift some Mediterranean ecosystems from annual C sinks to sources.

Comparison of Trends in Habitat and Resource Selection by the Spanish Festoon,Zerynthia rumina, and the Whole Butterfly Community in a Semi-Arid Mediterranean Ecosystem

Butterfly community and single species based approaches were taken to establish conservation priorities within a nature reserve in Central Spain. In this study, patch type (sclerophyllous, halophilous, or disturbed), potential herbaceous nectar availability, potential woody plant nectar availability, total nectar availability, and two approximations to plant diversity (herbaceous and woody plant diversity) were evaluated as variables that account for adult butterfly density. Butterfly communities in the reserve, which consist mostly of generalist species, were denser in relatively wet areas dominated by halophilous vegetation. Diversity did not significantly vary between ecologically different transects. Total nectar availability correlated with higher butterfly densities within both undisturbed and disturbed areas, which could be primarily explained by the lack of water typical of semi-arid Mediterranean climates, where fresh, nectariferous vegetation is scarce. Woody plants were also found to be important sources of nectar and shelter. In the dryer sclerophyllous sites, adult butterfly density was best explained by herbaceous plant diversity, suggesting better quality of available resources. The endangered specialist Zerynthia rumina (L.) (Lepidoptera: Papilionidae) was only present at the sclerophyllous sites. Its density was very low in all sampled transects, excluding one relatively isolated transect with high larval host-plant density. In contrast to the community-based approach, density of Z. rumina adults is better explained by the density of its larval host-plant than by nectar availability, a trend previously described for other sedentary species. Management strategies for protecting insect-rich areas should consider the specific ecological requirements of endangered species.

Simulated climate change reduced the capacity of lichen-dominated biocrusts to act as carbon sinks in two semi-arid Mediterranean ecosystems

The importance of biological soil crusts (biocrusts) for the biogeochemistry of drylands is widely recognized. However, there are significant gaps in our knowledge about how climate change will affect these organisms and the processes depending on them. We conducted a manipulative full factorial experiment in two representative dryland ecosystems from central (Aranjuez) and southeastern (Sorbas) Spain to evaluate how precipitation, temperature and biocrust cover affected the assimilation and net C balance of biocrusts. Chlorophyll fluorescence, net photosynthesis and dark respiration were measured in situ bimonthly during a year. We also conducted daily cycle measurements of net photosynthesis in winter and at the end of spring. In Sorbas, warming reduced the fixation of atmospheric C in biocrust-dominated microsites throughout the year. In Aranjuez, there was an interaction between the three factors evaluated; during winter, net photosynthesis was significantly greater in high biocrust cover plots under natural conditions and in the rainfall exclusion treatment. During the daily surveys, rainfall exclusion and warming reduced C fixation in Sorbas and in Aranjuez respectively. The effects of the treatments evaluated varied with the rainfall and non-rainfall water inputs (NRWIs) registered before the measurements. Our results suggest that changes in NRWI regimes as consequence of warming could have a greater impact on the C balance of biocrusts than changes in rainfall amounts. They also indicate that climate change may reduce the photosynthetic ability of lichens, with a consequent reduction of their dominance in biocrust communities at the mid to long term. This could reduce the ability of dryland ecosystems to fix atmospheric C.

Comparison of trends in habitat and resource selection by the Spanish Festoon, Zerynthia rumina, and the whole butterfly community in a semiarid Mediterranean ecosystem.

Butterfly community and single species based approaches were taken to establish conservation priorities within a nature reserve in Central Spain. In this study, patch type (sclerophyllous, halophilous, or disturbed), potential herbaceous nectar availability, potential woody plant nectar availability, total nectar availability, and two approximations to plant diversity (herbaceous and woody plant diversity) were evaluated as variables that account for adult butterfly density. Butterfly communities in the reserve, which consist mostly of generalist species, were denser in relatively wet areas dominated by halophilous vegetation. Diversity did not significantly vary between ecologically different transects. Total nectar availability correlated with higher butterfly densities within both undisturbed and disturbed areas, which could be primarily explained by the lack of water typical of semiarid Mediterranean climates, where fresh, nectariferous vegetation is scarce. Woody plants were also found to be important sources of nectar and shelter. In the dryer sclerophyllous sites, adult butterfly density was best explained by herbaceous plant diversity, suggesting better quality of available resources. The endangered specialist Zerynthia rumina (L.) (Lepidoptera: Papilionidae) was only present at the sclerophyllous sites. Its density was very low in all sampled transects, excluding one relatively isolated transect with high larval hostplant density. In contrast to the community-based approach, density of Z. rumina adults is better explained by the density of its larval hostplant than by nectar availability, a trend previously described for other sedentary species. Management strategies for protecting insect-rich areas should consider the specific ecological requirements of endangered species.

Retama-rhizobia symbiosis studies in some countries of the Mediterranean Basin

The bacteria called rhizobia are nitrogen-fixing bacteria which establish symbioses with plant species in the legume family. They induce positive interaction between soil microorganisms and the plant and are essential for plant growth in semiarid Mediterranean ecosystems that usually present severe water stress, scarcity of plants, low available nutrients and microbiological activity. Considered as one of the endemic legume shrubs of the Mediterranean basin, Retama plays an important role in nitrogen cycling, and thus, can be used to restore heavily disturbed and degraded soils. These species are adapted to harsh environmental conditions and their establishment and growth are enhanced by their ability to enter into effective symbiosis with multiple rhizobia species. In this article, we review the cumulative evidence of the Retama-rhizobia symbiosis in the Mediterranean basin and its ecological impact on revegetation and soil fertilization.

Simulated nitrogen deposition affects soil fauna from a semiarid Mediterranean ecosystem in central Spain

Nitrogen (N) deposition is a major threat to the semiarid Mediterranean ecosystems. We simulated a gradient of N deposition (0, 10, 20 and 50 kg N ha−1 year−1 + 6.4 kg N ha−1 year−1 ambient deposition) in a Mediterranean shrubland from central Spain. In autumn 2011 (after 4 years of experimental duration), soil cores were taken to extract the soil fauna. Acari (45.54%) and Collembola (44.00%) were the most represented taxonomical groups, and their abundance was negatively related to soil pH. Simulated N deposition had an impact on the total number of individuals in soil as well as on Collembola and Pauropoda abundance. Collembola abundance increased with N loads up to 20 kg N ha-1 year-1 and then decreased. This response was attributed to soil acidification (between 0 and 20 kg N ha-1 year-1) and increased soil ammonium (between 20 and 50 kg N ha-1 year-1). Pauropoda were favoured by additions of 50 kg N ha-1 year-1, and it was the only taxonomical group whose abundance was exclusively related to N deposition, suggesting their potential as bioindicators. Contrary to predictions, there was a negative relationship between soil faunal abundance and plant diversity. In conclusion, soil faunal communities from semiarid Mediterranean ecosystems in central Spain seem to be primarily influenced by soil chemistry (mainly pH) but are also susceptible to increased N deposition. The main drivers of change under increased N deposition scenarios seem to be soil acidification and increased ammonium in soils where nitrate is the dominant mineral N form.

On the importance of shrub encroachment by sprouters, climate, species richness and anthropic factors for ecosystem multifunctionality in semi-arid Mediterranean ecosystems.

One of the most important changes taking place in drylands worldwide is the increase of the cover and dominance of shrubs in areas formerly devoid of them (shrub encroachment). A large body of research has evaluated the causes and consequences of shrub encroachment for both ecosystem structure and functioning. However, there are virtually no studies evaluating how shrub encroachment affects the ability of ecosystems to maintain multiple functions and services simultaneously (multifunctionality). We aimed to do so by gathering data from ten ecosystem functions linked to the maintenance of primary production and nutrient cycling and storage (organic C, activity of β-glucosidase, pentoses, hexoses, total N, total available N, amino acids, proteins, available inorganic P and phosphatase activity), and summarizing them in a multifunctionality index (M). We assessed how climate, species richness, anthropic factors (distance to the nearest town, sandy and asphalted road, and human population in the nearest town at several historical periods) and encroachment by sprouting shrubs impacted both the functions in isolation and M along a regional (ca. 350 km) gradient in Mediterranean grasslands and shrublands dominated by a non-sprouting shrub. Values of M were higher in those grasslands and shrublands containing sprouting shrubs (43% and 62%, respectively). A similar response was found when analyzing the different functions in isolation, as encroachment by sprouting shrubs increased functions by 2%-80% compared to unencroached areas. Encroachment was the main driver of changes in M along the regional gradient evaluated, followed by anthropic factors and species richness. Climate had little effects on M in comparison to the other factors studied. Similar responses were observed when evaluating the functions in isolation. Overall, our results showed that M was higher at sites with higher sprouting shrub cover, longer distance to roads and higher perennial plant species richness. Our study is the first documenting that ecosystem multifunctionality in shrublands is enhanced by encroaching shrubs differing in size and leaf attributes. Our findings reinforce the idea that encroachment effects on ecosystem functioning cannot be generalized, and that are largely dependent on the traits of the encroaching shrub relative to those of the species being replaced.

Nitrogen deposition alters nitrogen cycling and reduces soil carbon content in low-productivity semiarid Mediterranean ecosystems

Anthropogenic N deposition poses a threat to European Mediterranean ecosystems. We combined data from an extant N deposition gradient (4.3–7.3 kg N ha−1 yr−1) from semiarid areas of Spain and a field experiment in central Spain to evaluate N deposition effects on soil fertility, function and cyanobacteria community. Soil organic N did not increase along the extant gradient. Nitrogen fixation decreased along existing and experimental N deposition gradients, a result possibly related to compositional shifts in soil cyanobacteria community. Net ammonification and nitrification (which dominated N-mineralization) were reduced and increased, respectively, by N fertilization, suggesting alterations in the N cycle. Soil organic C content, C:N ratios and the activity of β-glucosidase decreased along the extant gradient in most locations. Our results suggest that semiarid soils in low-productivity sites are unable to store additional N inputs, and that are also unable to mitigate increasing C emissions when experiencing increased N deposition.

Relationships Between Plant Spatial Patterns, Water Infiltration Capacity, and Plant Community Composition in Semi-arid Mediterranean Ecosystems Along Stress Gradients

Water redistribution from bare soil to vegetation patches is a key feature of semi-arid ecosystems, and is responsible for their patchy vegetation patterns. The magnitude of water redistribution depends on the properties of the bare soil (which determine the amount of water run-off) and the capacity of vegetation patches to trap water run-on. We examined the relationships between plant spatial patterns, water infiltration into bare soil, and plant community composition in semi-arid sites with different hydro-physical properties (silty and gypseous soils) in NE Spain. We also studied the effect of two stressors, aridity and grazing, on water infiltration and plant spatial patterns. Our results indicate a negative correlation of bare soil sorptivity (the capacity to absorb water by capillarity) and vegetation aggregation. There was a strong positive correlation between perennial grass cover and the spatial aggregation of vegetation, but aggregation was not associated with positive associations of different plant types. The aggregation of vegetation was positively correlated with species richness and the overall extent of vegetation cover. Grazing reduced water infiltration into silty soils, which are prone to compaction. In contrast, soil crust affected the hydrology of gypseous soils, especially in the most arid sites, where grazing increased infiltration, reducing surface sealing due to breaking of the soil crust. Together, our results suggest that biotic and abiotic factors affect the hydro-physical properties of soils in the semi-arid ecosystems of NE Spain, which is linked to the plant communities through the spatial distribution of plants.

Factors affecting vegetation and soil recovery in the Mediterranean woodland of the Canary Islands (Spain)

We investigated whether human-induced soil degradation may hinder the regeneration of a semiarid Mediterranean-type ecosystem with high biodiversity and conservation interest on the Canary Islands. To further this aim, the replacement of plant species and life-forms and the variation in soil quality were studied during the process of succession in old fields abandoned at different times and in relicts of the original thermophilous woodland by means of multivariate techniques of ordination (RDA, DCA, CCA) and clustering (k-means + IndVal). The studied ecosystem showed a limited capacity for recovery, which appears to be lower than in other similar semiarid Mediterranean ecosystems. Soil parent material exerted a considerable influence on plant reestablishment, and this process was more efficient with pumice-type rocks. Human activities have given rise to a loss of soil organic matter and an enrichment of certain nutrients, which revert to their normal levels once disturbance ceases. However, the erosion resulting from ploughing, farming and subsequent land abandonment has resulted in irreversible degradation of the soil water regime, thus severely limiting the restoration of the original ecosystem. We concluded that aridification due to soil degradation may, in certain cases, prevent the complete regeneration of thermophilous woodlands of the Canary Islands, and it must be taken into account when performing restoration activities. Pumice mulch can be useful for restoration, as it provides a suitable substrate in key stages of the plant recolonisation process.

Impact of burning and fertilization on dry Mediterranean grassland productivity and diversity

Fire alters plant community structure by changing dominant plant species and shifting resource mobilization, while fertilizer application alters plant community structure through its impact on productivity. Here we studied prescribed burning and fertilizer treatments on the productivity and diversity of a semi-arid Mediterranean grassland ecosystem. Variables measured were aboveground plant productivity, vegetation, litter and soil cover, species richness and plant density of an Avena/Hordeum (A-H) dominated plant community using a split-plot experimental design over a three-year period. Burning increased vegetative cover by 21.2% and reduced litter cover by 22.6%, while fertilization increased vegetative cover by 31.6%. The combined effect of burning and fertilization increased vegetative cover by 35.5% and reduced litter cover by 13.9%. Aboveground biomass was also higher in burned treatments (52.41 g 0.25 m−2) by 39.5% over unburned treatments. Burning reduced the proportion of the dominant Avena/Hordeum complex by 43.1%, thus significantly enhancing overall species richness by 52.5% over the unburned plots. On the other hand, fertilizer application increased biomass production and species richness by 37.4% and 13.5%, respectively, over the unfertilized plots. The combined effect of burning and fertilization enhanced biomass production and species richness by 90.8% and 69.8%, respectively, and reduced the dominant A-H by 45.5%. Burning generally resulted in lowering the dominance of the A-H complex compared to the control while fertilizer application had variable effect, reducing the proportion of A-H with burning but increasing it without burning. Reduction in the A-H complex increased the presence of other species, therefore enhancing diversity. Our findings showed that burning and fertilization treatments have a synergistic effect on both productivity and diversity of an arid grassland.

Analyzing the community composition of arbuscular mycorrhizal fungi colonizing the roots of representative shrubland species in a Mediterranean ecosystem

Community composition of arbuscular mycorrhizal (AM) fungi was analyzed in the roots of five representative shrub species (Genista cinerea, Lavandula latifolia, Rosmarinus officinalis, Thymus mastichina and Thymus zygis) growing in a typical semi-arid Mediterranean ecosystem. Roots from a well-preserved area of the ecosystem were extracted from soil and analyzed by nested PCR, single strand conformation polymorphism and sequencing of the NS31-AM1 and NS8-ARCH1311 regions of the small subunit of the ribosomal DNA (SSU rDNA). Ten AM fungal phylotypes were identified; eight belonged to the Glomeraceae and two to the Diversisporaceae. Only two of the phylotypes clustered with sequences of morphologically defined species and a high dominance by one AM group (Glomus intraradices) was detected. Our diversity analyses revealed that the AM fungal communities of G. cinerea, L. latifolia and T. mastichina did not significantly differ while the AM fungal communities of R. officinalis and T. zygis were distant from this cluster and from each other. The highest diversity was found in the roots of T. zygis. Our data indicate that co-occurring plant species may house distinct communities of AM fungi.

Diversity and Activity of Denitrifiers of Chilean Arid Soil Ecosystems

The Chilean sclerophyllous matorral is a Mediterranean semiarid ecosystem affected by erosion, with low soil fertility, and limited by nitrogen. However, limitation of resources is even more severe for desert soils such as from the Atacama Desert, one of the most extreme arid deserts on Earth. Topsoil organic matter, nitrogen and moisture content were significantly higher in the semiarid soil compared to the desert soil. Although the most significant loss of biologically preferred nitrogen from terrestrial ecosystems occurs via denitrification, virtually nothing is known on the activity and composition of denitrifier communities thriving in arid soils. In this study we explored denitrifier communities from two soils with profoundly distinct edaphic factors. While denitrification activity in the desert soil was below detection limit, the semiarid soil sustained denitrification activity. To elucidate the genetic potential of the soils to sustain denitrification processes we performed community analysis of denitrifiers based on nitrite reductase (nirK and nirS) genes as functional marker genes for this physiological group. Presence of nirK-type denitrifiers in both soils was demonstrated but failure to amplify nirS from the desert soil suggests very low abundance of nirS-type denitrifiers shedding light on the lack of denitrification activity. Phylogenetic analysis showed a very low diversity of nirK with only three distinct genotypes in the desert soil which conditions presumably exert a high selection pressure. While nirK diversity was also limited to only few, albeit distinct genotypes, the semiarid matorral soil showed a surprisingly broad genetic variability of the nirS gene. The Chilean matorral is a shrub land plant community which form vegetational patches stabilizing the soil and increasing its nitrogen and carbon content. These islands of fertility may sustain the development and activity of the overall microbial community and of denitrifiers in particular.

Stand structure modulates the long‐term vulnerability ofPinus halepensisto climatic drought in a semiarid Mediterranean ecosystem

We investigated whether stand structure modulates the long-term physiological performance and growth of Pinus halepensis Mill. in a semiarid Mediterranean ecosystem. Tree radial growth and carbon and oxygen stable isotope composition of latewood (δ(13)C(LW) and δ(18)O(LW), respectively) from 1967 to 2007 were measured in P. halepensis trees from two sharply contrasting stand types: open woodlands with widely scattered trees versus dense afforested stands. In both stand types, tree radial growth, δ(13)C(LW) and δ(18)O(LW) were strongly correlated with annual rainfall, thus indicating that tree performance in this semiarid environment is largely determined by inter-annual changes in water availability. However, trees in dense afforested stands showed consistently higher δ(18)O(LW) and similar δ(13)C(LW) values compared with those in neighbouring open woodlands, indicating lower stomatal conductance and photosynthesis rates in the former, but little difference in water use efficiency between stand types. Trees in dense afforested stands were more water stressed and showed lower radial growth, overall suggesting greater vulnerability to drought and climate aridification compared with trees in open woodlands. In this semiarid ecosystem, the negative impacts of intense inter-tree competition for water on P. halepensis performance clearly outweigh potential benefits derived from enhanced infiltration and reduced run-off losses in dense afforested stands.

Temporal dynamics of arbuscular mycorrhizal fungi colonizing roots of representative shrub species in a semi-arid Mediterranean ecosystem

Arbuscular mycorrhizal (AM) symbiosis plays an important role in improving plant fitness and soil quality, particularly in fragile and stressed environments, as those in certain areas of Mediterranean ecosystems. AM fungal communities are usually affected by dynamic factors such as the plant community structure and composition, which in turn are imposed by seasonality. For this reason, a one-year-round time-course trial was performed by sampling the root system of two representative shrubland species (Rosmarinus officinalis and Thymus zygis) within a typical Mediterranean ecosystem from the Southeast of Spain. The 18S rDNA gene, of the AM fungal community in roots, was subjected to PCR-SSCP, sequencing, and phylogenetic analysis. Forty-three different AM fungal sequence types were found which clustered in 16 phylotypes: 14 belonged to the Glomeraceae and two to the Diversisporaceae. Surprisingly, only two of these phylotypes were related with sequences of morphologically defined species: Glomus intraradices and Glomus constrictum. Significant differences were detected for the relative abundance of some phylotypes while no effects were found for the calculated diversity indices. These results may help to design efficient mycorrhizal-based revegetation programs for this type of ecosystems.

Habitat selection by European badgers in Mediterranean semi-arid ecosystems

We studied the habitat selection patterns of badgers Meles meles (Linnaeus, 1758) in Mediterranean semi-arid ecosystems. Fifty-seven plots were sampled in two semi-arid regions of Spain. In each plot, badger latrines were located along 2.6 km transects. The number of badger latrines per km was used as a surrogate of badger abundance and as an index of habitat selection by badgers. For each plot, a series of environmental variables were measured at two spatial scales. These variables were related to land use and vegetation formation parameters that are considered potentially important for habitat requirements (i.e., food and shelter). The habitat selection model was carried out using generalised linear models (GLM) and an information-theoretic approach. Results indicated that badgers prefer fruit orchards, and shrub and rock-covered areas, which provide additional trophic and shelter resources, and avoid intensively cultivated fields and human settlements. We conclude that badger conservation in semi-arid environments of the Iberian Peninsula requires the existence of fruit orchards and the limitation of human development. Policies restraining agriculture intensification would encourage traditional or new non-intensive agricultural practices and increase shrub-patch availability, which would benefit this species.

Do biotic interactions modulate ecosystem functioning along stress gradients? Insights from semi-arid plant and biological soil crust communities.

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative-competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant-plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.

Scale-dependent effects of pines on the herbaceous layer diversity in a semi-arid mediterranean ecosystem

In arid and semi-arid ecosystems, the presence of woody neighbours affects the existence of several herbaceous species by modifying critical aspects of the environment (e.g., soil humidity, nutrient content or light availability) beneath their canopies. Herbaceous species growing in the understory of Pinus pinaster may be distinct from those in open areas due to litter fall, light interception and changes in nutrient availability. We suggest that the overall effect of woody neighbours on herbaceous layer diversity may vary with the scale focus of analysis. To examine this hypothesis, we collected data on the abundance of herbaceous species in open pineland forests of the central Iberian Peninsula (Spain) using sample quadrats of 0.5 m × 0.5 m distributed beneath, at the edge, and outside the canopy of pines in a landscape composed of dunes and plains. The results of CCA ordination revealed significant spatial segregation of herbaceous species reflecting the occurrence of pines and dunes in the landscape. ...

Nitrogen fertilization and water supply affect germination and plant establishment of the soil seed bank present in a semi-arid Mediterranean scrubland

Anthropogenic nitrogen (N) inputs in terrestrial ecosystems are higher than those that occur naturally and have been related to global biodiversity loss and altered ecosystem functioning. However, its effects on Mediterranean-type ecosystems, where production is water-limited and N regulated, remain unclear. We conducted a green-house experiment where we evaluated the effects of four simulated scenarios of N pollution (0, 10, 20 and 50 kg N ha−1 year−1) and two differential water supply regimes on the germination (experiment 1) and early plant establishment (experiment 2) of a seed bank from a semi-arid Mediterranean ecosystem of central Spain. Seed bank density was estimated as 62,374 ± 3,279 seeds m−2. Approximately 99.5% of emerged seeds corresponded to only 14 species of a total of 52, the majority of which were the annual forb Sagina apetala. The responses for N treatments were species-specific, mainly positive or unimodal, with watering treatments having some interactive effects. N and water supply also affected total and specific productivity; the responses found for N treatments were mainly humpback-shaped and an increased water supply had additive effects on community establishment in terms of total plant biomass. This response was linked to forb responsiveness. Contrary to predictions, grass biomass did not change with N supply; however, grass to forb ratio was affected because of changes in the latter. Overall, these experiments suggest a critical load for plant biomass production and conclude that N and water availability and supply can modify germination and plant establishment. This should be taken into account when analysing the effects of global change on the dynamics of plant communities where annuals are dominant or vegetation must establish from seed following a natural or anthropogenic disturbance regime.