Retama Sphaerocarpa Research Articles

An Indigenous Drought-Tolerant Strain of Glomus intraradices Associated with a Native Bacterium Improves Water Transport and Root Development in Retama sphaerocarpa

The effects of interactions between Bacillus thuringiensis, a drought-adapted bacterium, and two isolates of Glomus intraradices, an arbuscular mycorrhizal (AM) fungus, on Retama sphaerocarpa, a drought-adapted legume, were investigated. The fungal isolates were an indigenous drought-tolerant and a nonindigenous drought-sensitive isolate. Shoot length and root growth, symbiotic parameters, water transport (in terms of percent relative plant water uptake), and volumetric soil moisture and soil enzymatic activities in response to microbial inoculations were evaluated. Retama plants colonized by G. intraradices plus Bacillus possessed similar shoot length after 30 days from sowing compared with noninoculated Retama plants after 150 days. Inoculation with drought-adapted bacterium increased root growth by 201%, but maximum root development was obtained by co-inoculation of B. thuringiensis and the indigenous G. intraradices. Nodules were formed only in plants colonized by autochthonous AM fungi. Relative water uptake was higher in inoculated than in noninoculated Retama plants, and these inoculants depleted soil water content concomitantly. G. intraradices-colonized Retama reached similar shoot length irrespective of the fungal origin, but there were strong differences in relative water uptake by plants colonized by each one of the fungi. Indigenous G. intraradices-colonized roots (evaluated as functional alkaline phosphatase staining) showed the highest intensity and arbuscule richness when associated with B. thuringiensis. The interactive microbial effects on Retama plants were more relevant when indigenous microorganisms were involved. Co-inoculation of autochthonous microorganisms reduced by 42% the water required to produce 1 mg of shoot biomass. This is the first evidence of the effectiveness of rhizosphere bacterium, singly or associated with AM fungus, in increasing plant water uptake, which represents a positive microbial effect on plants grown under drought environments.

Exposure and bioavailability of arsenic in contaminated soils from the La Parrilla mine, Spain

Arsenic derived from mining activity may contaminate water, soil and plant ecosystems resulting in human health and ecotoxicological risks. In this study, exposure assessment of arsenic (As) in soil, spoil, pondwater and plants collected from the areas contaminated by mine tailings and spoils in and around the La Parrilla mine, Caceres province, Spain, was carried out using AAS method. Water solubility, bioavailability and soil–plant transfer coefficients of As and phytoremediation potential of plants were determined. Arsenic concentrations varied from 148 to 2,540 mg/kg in soils of site 1 and from 610 to 1,285 mg/kg in site 2 exceeding the guideline limit for agricultural soil (50 mg/kg). Arsenic concentrations in pond waters varied from 8.8 to 101.4 μg/l. High concentrations of water-soluble As in the soils that ranged from 0.10 to 4.71 mg/kg in site 1 and from 0.46 to 4.75 mg/kg in site 2 exceeded the maximum permitted level of water-soluble As (0.04 mg/kg) in agricultural soils. Arsenic concentrations varied from 0.8 to 149.5 mg/kg dry wt in the plants of site 1 and from 2.0 to 10.0 mg/kg in the plants of site 2. Arsenic concentrations in plants increased in the approximate order: Retama sphaerocarpa < Pteridium aquilinum < Erica australis < Juncus effusus < Phalaris caerulescens < Spergula arvensis in site 1. The soil–plant transfer coefficients for As ranged from 0.001 to 0.21 in site 1 and from 0.004 to 0.016 in site 2. The bioconcentration factor based on water-soluble As of soil varied from 3.2 to 593.9 in the plants of site 1 whereas it varied from 2.1 to 20.7 in the plants of site 2. To our knowledge, this is the first study in Europe to report that the fern species P. aquilinum accumulates extremely low contents of As in its fronds despite high As levels in the soils. Therefore, the S. arvensis, P. caerulescens and J. effusus plant species grown in this area might be used to partly remove the bioavailable toxic As for the purpose of minimization of mining impacts until hypothetical hyperaccumulating and/or transgenic plants could be transplanted for the phytoremediation of As contaminated soils.

Shrubs as a source of spatial heterogeneity—the case of Retama sphaerocarpa in Mediterranean pastures of central Spain

The processes underlying the positive effect exerted by woody species on other plant species can be grouped into a physical effect due to the canopy itself (mainly amelioration of temperature and light extremes), and an edaphic effect (nutrient enrichment). Both groups of effects are present in the savanna-like systems originated by Retama sphaerocarpa in the Iberian Peninsula. The consequences of these shrubs for the spatial heterogeneity induced in the herbaceous community are not well known. The herbaceous community was sampled by means of radial transects around 20 adult shrubs, from the canopy centre to the open areas among the shrubs, both northwards and southwards. Floristic composition, diversity and pasture mean height change both along the inside–outside gradient, and from north to south. As a nutrient enriched environment, the centre of the understorey showed the lowest species richness and evenness, and the highest standing crop. In contrast, the environmentally stressful open areas showed the highest number of species and evenness, and the lowest standing crop. This general effect is only partially due to nutrient availability, and is not homogeneous around the shrub, as open area conditions penetrate into the understorey driving the herbaceous community towards that of open areas. An individual shrub effect is also suggested by our results. Different conditions are thus provided by R. sphaerocarpa understoreys, favouring different subsets of species from the general pool, and therefore defining shrub canopies as sources of spatial heterogeneity in the whole savanna-like system.

Interaction between AM fungi and a liquid organic amendment with respect to enhancement of the performance of the leguminous shrub Retama sphaerocarpa

This study examined the interactions between the inoculation with three arbuscular mycorrhizal fungi, namely, Glomus intraradices, Glomus deserticola and Glomus mosseae, and the addition of a liquid organic amendment at different rates (0, 50, 100 or 300 mg C of liquid amendment per kilogram soil) obtained by alkaline extraction of composted dry olive residue with respect to their effects on growth of Retama sphaerocarpa seedlings and on some microbiological and physical properties of soil. One year after planting, both mycorrhizal inoculation treatments and the addition of amendment had increased plant growth and dehydrogenase, urease and benzoyl argininamide hydrolysing activities. The inoculation with G. mosseae increased plant growth to a greater extent than the addition of the amendment (about 35% greater than plants grown in the amended soil and about 79% greater than control plants) and both treatments produced similar increases in soil aggregate stability (about 31% higher than control soil). The organic amendment produced a very significant decrease in the levels of microbial biomass C and a strong increase in soil dehydrogenase and urease activities, which were proportional to the amendment rate. Only the combined treatment involving the addition of a medium dose of amendment (100 mg C kg−1 soil) and the mycorrhizal inoculation with G. intraradices or G. deserticola produced an additive effect on the plant growth with respect to the treatments applied individually (about 77% greater than plants grown in the amended soil and about 63% greater than inoculated plants).

Colonization processes in semi-arid Mediterranean old-fields

Semi-arid Mediterranean ecosystems undergo a slow process of secondary succession after disturbance which is dependent on rainfall. We tested the hypothesis that colonization of abandoned lands in semi-arid SE Spain was slowed down by seed dispersal and species-specific responses to climatic variability. In field and glasshouse experiments we analysed germination of the two main colonizer species, their appearance in the field as seedlings, their relationships with climate and the resulting population structure. Seed germination of the two dominant legume species, Anthyllis cytisoides and Retama sphaerocarpa, was enhanced by scarification, but triggered in the field by temperature and rainfall, respectively. There was no evidence of allelopathic effects affecting germination in the field. Population structure reflected high inter-annual variability in shrub establishment, which was related to rainfall in spite of self-thinning and other processes that modeled current shrub numbers. Colonization of abandoned lands in semi-arid, Mediterranean environments in SE Spain seems to be primarily limited by seed dispersal, but also affected by climatic variability, which produced temporally separated recruitment events.

Nutrient acquisition and nitrate reductase activity of mycorrhizal &lt;I&gt;Retama sphaerocarpa&lt;/I&gt; L. seedlings afforested in an amended semiarid soil under two water regimes

Nutrient acquisition and nitrate reductase activity of mycorrhizal &lt;I&gt;Retama sphaerocarpa&lt;/I&gt; L. seedlings afforested in an amended semiarid soil under two water regimes

Sheep gut passage and survival of Mediterranean shrub seeds

Although viable seeds of Mediterranean dry-fruited shrubs are found in herbivore dung, the ecological importance of this observation is still not well understood. We analysed seed retrieval percentages, defecation time and germinability after sheep gut passage for the five most common shrub species of an area in central Spain (Retama sphaerocarpa, Cytisus scoparius, Halimium umbellatum subsp. viscosum, Cistus ladanifer and Lavandula stoechas subsp. pedunculata). Five ewes were fed seeds, and their dung was collected regularly during the following week. Seeds were hand-sorted from dung subsamples and tested for germinability. The defecated seeds were clustered in time, with a majority retrieved in the 24–40 h period, although over 1% of the seeds were retained in the gut for more than 72 h. Data suggested a possible link between seed size and retrieval, with medium-sized seeds less damaged (16–23%) than larger and smaller seeds (10–12%), although only a small number of species were studied. Germination results showed an increased percentage of germination after gut passage for H. umbellatum (x2 test, P&lt;0.05) and a marginally significant difference for C. scoparius (P&lt;0.1). Soft-seeded L. stoechas did not germinate after gut passage. The results indicate a potential role of herbivore endozoochory for the long-distance dispersal of dry-fruited shrubs and their potential colonization of distant sites.

Nutrient acquisition and nitrate reductase activity of mycorrhizal Retama sphaerocarpa L. seedlings afforested in an amended semiarid soil under two water regimes

Abstract. We studied the effect of inoculation with three arbuscular‐mycorrhizal (AM) fungi (Glomus intraradices Schenck &amp; Smith, Glomus deserticola (Trappe, Bloss. &amp; Menge) and Glomus mosseae ([Nicol &amp; Gerd.] Gerd. &amp; Trappe) and the addition of composted sewage sludge on root nitrate reductase (NR, EC 1.6.6.1.) activity, mycorrhizal colonization, plant growth and nutrient uptake in Retama sphaerocarpa L. seedlings afforested in a semiarid, degraded Mediterranean soil under well‐watered and non‐watered conditions. Six months after planting, the mycorrhizal inoculation and the irrigation of plants had a strong effect on the growth parameters. The effect on plant growth was a negative interaction between plant irrigation and mycorrhizal inoculation and a positive interaction between plant irrigation and composted sewage sludge addition. The latter treatment had a significant, but moderate, effect on the growth but conferred no additional benefit when combined with mycorrhizal inoculation. Mycorrhizal inoculation, composted sewage sludge and irrigation had a significant effect on NR activity in roots and on foliar nutrients. The irrigation significantly increased the positive effect of composted sewage sludge on NR activity and the concentrations of foliar N and K. The effect of mycorrhizal inoculation on NR activity did not depend on the water regime. The effectiveness of mycorrhizal inoculation on the establishment and growth of R. sphaerocarpa seedlings in these Mediterranean conditions was independent of water regime. The addition of composted sewage sludge was only effective when soil water was freely available. The combination of mycorrhizal inoculation and composted sewage sludge addition had no synergistic effect on plant growth.

Changes in rhizosphere microbial activity mediated by native or allochthonous AM fungi in the reafforestation of a Mediterranean degraded environment

This study was carried out in a semiarid degraded area to assess the effectiveness of mycorrhizal inoculation with a mixture of native arbuscular mycorrhizal (AM) fungi or an allochthonous AM fungus (Glomus claroideum), on the establishment of Olea europaea subsp. sylvestris L. and Retama sphaerocarpa (L.) Boissier in this area. Associated changes in the soil microbiological properties and aggregate stability related to these AM inocula were also recorded. Eighteen months after planting, G. claroideum had increased available P in the rhizosphere of both shrub species. In general, both inoculation treatments increased water-soluble C and water-soluble and total carbohydrates, G. claroideum being the most effective inoculum, particularly in R. sphaerocarpa. The mixture of native AM fungi was the most effective treatment for increasing the aggregate stability of R. sphaerocarpa soil, while that of O. europaea was increased only by G. claroideum. Increased (dehydrogenase, urease, protease-BAA, acid phosphatase and β-glucosidase) enzyme activities, in particular of dehydrogenase and acid phosphatase, were recorded in the rhizosphere of both mycorrhizal shrub species. The mixture of native AM fungi was the most effective treatment for stimulating the growth of O. europaea and R. sphaerocarpa (11.6-fold and 3.3-fold, respectively, greater than control plants). The establishment of mycorrhizal shrub species favoured the reactivation of soil microbial activity, which was linked to an increase in aggregate stability.

Developmental instability and plant potential fitness in a Mediterranean perennial plant, Retama sphaerocarpa (L.) Boiss

Developmental instability is manifested as developmental errors reflected in exaggerated intra-individual variation in repeated traits and patterns. Plants, as organisms with modular construction, are very suitable subjects for detecting developmental instability caused by environmental disturbance. The analysis of the asymmetry of plant structural traits allows for determination of deviations from the basic structural pattern, which is a measure of plant developmental instability. In this paper, we study the relationship between intra-individual variations on self-similar structural traits (as a measure of developmental instability) and plant potential fitness. Randomly-selected branches (composed of branch segments) were monitored on different plants of a natural population of the woody perennial plant Retama sphaerocarpa (L.) Boiss. Data on the morphology and the demographic processes that occurred during plant development (determined from marks left on the persistent structure of the plant) were recorded on the different branches. Different measures of developmental instability were analysed and related with plant potential fitness, which was estimated from the demographic data of the modules of each plant (bud development, branch survival, etc). Our results show a direct relationship between developmental instability measured on structural traits (except for branch segment diameter) and plant potential fitness, estimated by means of branch survival.

Establishment of Retama sphaerocarpa L. seedlings on a degraded semiarid soil as influenced by mycorrhizal inoculation and sewage‐sludge amendment

AbstractA field experiment was carried out to evaluate the effectiveness of mycorrhizal inoculation with three arbuscular mycorrhizal (AM) fungi (Glomus intraradices Schenck &amp; Smith, Glomus deserticola (Trappe, Bloss. &amp; Menge), and Glomus mosseae (Nicol &amp; Gerd.) Gerd. &amp; Trappe) and the addition of composted sewage sludge (SS) with respect to the establishment of Retama sphaerocarpa L. seedlings, in a semiarid Mediterranean area. Associated changes in soil chemical (nutrient content and labile carbon fractions), biochemical (enzyme activities), and physical (aggregate stability) parameters were observed. Six months after planting, both the addition of composted SS and the mycorrhizal‐inoculation treatments had increased total N content, available‐P content, and aggregate stability of the soil. Values of water‐soluble C and water‐soluble carbohydrates were increased only in the mycorrhizal‐inoculation treatments. Rhizosphere soil from the mycorrhizal‐inoculation treatments had significantly higher enzyme activities (dehydrogenase, protease‐BAA, acid phosphatase, and β‐glucosidase) than the control soil. In the short‐term, mycorrhizal inoculation with AM fungi was the most effective treatment for enhancement of shoot biomass, particularly with G. mosseae (about 146% higher with respect to control plants). The addition of the composted SS alone was sufficient to restore soil structural stability but was not effective with respect to improving the performance of R. sphaerocarpa plants.

Survival of inocula and native AM fungi species associated with shrubs in a degraded Mediterranean ecosystem

Reconstitution of the potential of soil mycorrhizal inoculum is a key step in revegetation programs for semiarid environments. We tested the effectiveness of inoculation with native arbuscular mycorrhizal (AM) fungi or with an allochthonous AM fungus, Glomus claroideum, with respect to the growth of four shrub species, the release of mycorrhizal propagules in soil, within and outside the canopy, and the improvement of soil structural stability. Two years after outplanting, the mixture of native endophytes was more effective than, for Olea europaea subsp. sylvestris, Retama sphaerocarpa and Rhamnus lycioides, or equally as effective as, for Pistacia lentiscus, the non-native AM fungus Glomus claroideum, with respect to increasing shoot biomass and foliar NPK contents. The increases in glomalin concentration and structural stability produced by inoculation treatments in the rhizosphere soil of the all shrub species, except R. lycioides, ranged from about 55 to 173% and 13 to 21%, respectively. The mixture of native AM fungi produced the highest levels of mycorrhizal propagules in soil from the center of the canopy of P. lentiscus and R. lycioides, while plants of O. europaea and R. sphaerocarpa inoculated with G. claroideum had more mycorrhizal propagules than did those inoculated with the mixture of native fungi. The number of mycorrhizal propagules in soil outside the canopy of the four shrub species was 5–35 times higher in inoculation treatments than in soil of the non-inoculated plants.

Is the interaction between Retama sphaerocarpa and its understorey herbaceous vegetation always reciprocally positive? Competition–facilitation shift during Retama establishment

Retama sphaerocarpa is a Mediterranean shrub that when adult, facilitates the establishment of herbaceous plants under its canopy. We test the hypothesis that during the establishment of R. sphaerocarpa seedlings, the interaction with the herbaceous plants is negative. We carried out a greenhouse experiment in which seedlings of R. sphaerocarpa were grown under different conditions of competition with herbs, watering and date of emergence. Measurements of seedling mortality, biomass and growth were taken during the first growing season. We found a significant relationship between R. sphaerocarpa seedling mortality and competition in early spring, presumably due to higher water demand of herbaceous plants. Generally, presence of herbaceous species, lower availability of water and late emergence had negative effects on biomass and growth of Retama seedlings. Additional water compensated for the negative effects of competition, except on leaves and cladodes of Retama seedlings, suggesting that other resources, such as light, could be the subject of competition. In contrast, Retama seedlings exerted a positive influence on the herbaceous plants by increasing their survival and biomass, probably as a consequence of the high availability of nutrients provided by the Rhizobia nodules in the roots of Retama seedlings. We concluded that, at the regeneration stage of the shrub, the interaction between the herbaceous vegetation and the shrub is negative for the shrub and positive for the herbs. This suggests a shift from competition to facilitation with age of Retama, as reciprocal positive interactions have been described between herbaceous plants and adult individuals of the shrub. © 2004 Elsevier SAS. All rights reserved.

Structural cost of shoot modules and its implications on plant potential fitness in a Mediterranean perennial shrub, Retama sphaerocarpa (L.) Boiss

This paper introduces a methodology to analyse the structural costs on plant potential fitness, empirically exemplified in the hierarchical shoot system of a Mediterranean perennial plant, Retama sphaerocarpa (L.) Boiss. During growing season every year (March–August), the terminal shoot (which is the basic unit of growth) develops inflorescences, flowers and fruits, as well as new shoots (first-, second- and third-order branching shoots) which have the potential to “behave” as terminal shoots in the following year. Different morphological and demographical aspects of the modules within the terminal shoot were measured in 100 terminal shoots selected from different plants of a natural population of R. sphaerocarpa. Complementary samples of 100 shoots of different branching orders were collected to obtain biomass estimations of the terminal shoots. We propose a simple procedure to estimate structural cost (biomass investment) on plant potential fitness (flowering buds) as a methodology for interpreting and comparing the consequences on fitness of different plant growth patterns. The results of this study exemplify how differential allocation patterns among plant structural modules, depending on their position within the shoot system, can be quantified to estimate their influence upon plant potential fitness.

Analysis of the legume-rhizobia symbiosis in shrubs from central western Spain.

This work analyses the diversity of rhizobia associated with some of the predominant shrubby legumes in central-western Spain. Symbiotic promiscuity and effectiveness were studied using cross-inoculation experiments with shrubby species. Six new bradyrhizobia strains were isolated from nodules collected from wild plants of six leguminous species, Cytisus balansae, C. multiflorus, C. scoparius, C. striatus, Genista hystrix and Retama sphaerocarpa. These isolates were genetically characterized by 16S rDNA partial sequencing and random amplification of polymorphic DNA-PCR fingerprinting. The phylogenetic analysis revealed that these isolates could represent three new Bradyrhizobium species. Shrubby legumes and bradyrhizobia displayed a high symbiotic promiscuity both for infectivity and effectiveness. Symbioses were effective in more than 70% of the associations established by four of the six plant species. Native woody legumes in western Spain are nodulated by Bradyrhizobium strains. The high degree of symbiotic promiscuity and effectiveness highlights the complex dynamics of these communities in wild ecosystems under a Mediterranean-type climate. Furthermore, the results from this study suggest a potential importance of inoculation for these legume species in soil-restoration projects. This is the first study, to our knowledge, that combines both molecular analysis and pot trials to study the rhizobia-legume symbiosis for wild legumes.

Water Relations Only Partly Explain the Distributions of Three Perennial Plant Species in a Semi-arid Environment

The water relations and stomatal conductances of three perennial plant species, Stipa tenacissima L., Anthyllis cytisoides L., and Retama sphaerocarpa (L.) Boiss., dominant on the upper slopes, mid-slopes and floor of a valley, respectively, in semi-arid south-east Spain, were investigated to test the hypothesis that differences in plant-soil water relations could account for the different distributions of each species in the catena. Diurnal measurements of water potential (Ψw), relative water content (RWC) and stomatal conductance (gs) of leaves were made over one year. Leaf temperature, air humidity, wind-speed and incident quantum flux density were measured concurrently. Soil water content was determined gravimetrically at 0 - 5 cm and 15 - 20 cm depths. Measurements of Ψw, RWC and gs were analysed according to meteorological conditions, based on the maxima for daily air temperature and atmospheric saturation water vapour deficit and on soil moisture content. The hypothesis that plant-soil water relations can explain the distribution of the three species along the catena from valley side to valley floor was rejected for Anthyllis and Stipa but confirmed for Retama.

Antioxidant enzyme activities in shoots from three mycorrhizal shrub species afforested in a degraded semi‐arid soil

Mycorrhizae may help plants to thrive in Mediterranean semi‐arid ecosystems by altering antioxidant enzyme activities. Our objective was to determine the influence of mycorrhizal inoculation with an allochthonous arbuscular mycorrhizal (AM) fungus, Glomus claroideum, Schenck &amp; Smith, or with a mixture of native AM fungi, on the activity of antioxidant enzymes from shoots of Olea europaea L. ssp. sylvestris, Retama sphaerocarpa (L.) Boissier and Rhamnus lycioides L. seedlings afforested in a degraded Mediterranean semi‐arid soil. One year after planting, shoot biomass of inoculated O. europaea seedlings was about 630%, of non‐inoculated plants. Shoot biomass of G. claroideum‐colonized R. sphaerocarpa was greater than that of seedlings inoculated with the mixed native AM fungi after 12 months. Inoculation with a mix of native AM fungi was the most effective treatment for increasing shoot biomass and N, P and K contents in shoot tissues of R. lycioides. Both mycorrhizal inoculation treatments increased the nutrient contents in shoots of O. europaea and R. lycioides. In O. europaea plants, the inoculation treatments increased catalase, ascorbate peroxidase and dehydroascorbate reductase activities, but not monodehydroascorbate reductase and glutathione reductase activities. Inoculation with G. claroideum increased the activities of all antioxidant enzymes in R. sphaerocarpa. Monodehydroascorbate reductase, glutathione reductase and superoxide dismutase activities in R. lycioides leaves were preferentially increased by inoculation with the mixture of native AM fungi. This work support the view that increased antioxidant enzyme activities could be involved, at least in part, in the beneficial effects of mycorrhizal colonization on the performance of shrub species grown under semi‐arid Mediterranean conditions.

Application of composted urban residue enhanced the performance of afforested shrub species in a degraded semiarid land

Improvement of physical–chemical soil quality is a key step for carrying out revegetation programs of degraded lands in Mediterranean semiarid areas. Organic residue addition may restore the quality of these areas. A field experiment was conducted in a silt-loam soil (Typic Petrocalcid) from a degraded semiarid Mediterranean area to evaluate the effect of the addition of a composted urban residue on soil aggregate stability, bulk density and chemical properties and on the establishment of Pistacia lentiscus and Retama sphaerocarpa seedlings. The composted residue was applied at a rate of 6.7 kg m −2 before planting. The nutrient content (NPK), total organic C and water soluble C were increased and bulk density was decreased, in the rhizosphere soil of both shrub species, by the composted residue. The addition of composted residue significantly increased the soil aggregate stability by about 22% for both shrub species. The beneficial effect of the composted residue on soil quality still persisted 18 months after addition. Eighteen months after planting, the addition of composted residue to soil had increased significantly the production of shoot biomass by P. lentiscus and R. sphaerocarpa, by about 160% and 320% respectively, compared to control values. Composted residue addition to soil can be considered an effective preparation method of a degraded area for carrying out successful revegetation programs with Mediterranean shrubs under semiarid conditions.

Soil fertility and herb facilitation mediated by Retama sphaerocarpa

Abstract. Woody legumes growing in dry climates can increase soil nutrient content and facilitate plant growth in their understorey. We investigated differences in soil fertility and herbaceous community in relation to the presence and absence of the legume Retama sphaerocarpa in a shrubland in a mediterranean type climate. The results showed a higher content of limiting nutrients for plant growth, such as phosphorus and nitrogen, under the R. sphaerocarpa canopy. Herbaceous biomass, cover and nitrogen content increased below the canopy. However, species richness and diversity were diminished in the presence of a dense canopy of R. sphaerocarpa. Nitrogen isotopic analysis did not indicate a clear relationship between symbiotic fixation in R. sphaerocarpa and nitrogen content of soil and plants under its canopy. Nevertheless, herbs growing in the understorey showed a linear correlation between foliar N content and 15N values. The existence of a dense shrub canopy induced a smaller monthly variation in herb richness, diversity and biomass, suggesting that it provides a stable microhabitat facilitating herbaceous vegetation establishment and growth. The ‘shelter’ effect was more important when environmental conditions became stressful for herbaceous plants.

Relationship between net radiation and solar radiation for semi-arid shrub-land

This paper presents the results obtained by analysing a set of measurements of surface radiation balance components at a semi-arid location in SE Spain. The relationships between net and surface shortwave radiation were explored by using 38 months of 5 min surface radiation. The study area is covered by sparse clumped shrub-land of different species, although close to the radiometric station Retama sphaerocarpa is dominant. After this study, it is evident that a single model based on surface shortwave radiation could provide appropriate estimates. The inclusion of albedo information provides a slight improvement on the estimation model. The use of seasonal models provides an appreciable improvement, while the benefits of including cloud information are only marginal.