Adrián Escudero

Plant species richness and ecosystem multifunctionality in global drylands

Global Ecosystem Analysis The relationship between species richness and the functional properties of their ecosystems has often been studied at small scales in experimental plots. Maestre et al. (p. 214; see the Perspective by Midgley) performed field measurements at 224 dryland sites from six continents and assessed 14 ecosystem functions related to carbon, nitrogen, and phosphorus cycling. Positive relationships were observed between perennial plant species richness and ecosystem functionality. The relative importance of biodiversity was found to be as large as, or larger than, many key abiotic variables. Thus, preservation of plant biodiversity is important to buffer negative effects of climate change and desertification in drylands, which collectively cover 41% of Earths land surface and support over 38% of the human population. Plant species richness is positively related to ecosystem multifunctionality in drylands at a global scale. Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth’s land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.

Spatial analysis of genetic diversity as a tool for plant conservation

Development of suitable approaches to the analysis of genetic diversity in a spatial context, where factors such as pollination, seed dispersal, breeding system, habitat heterogeneity and human influence are appropriately integrated, can provide new insights in the understanding of the mechanisms of maintenance and dynamics of populations. In this sense, it is important to recognise that patterns and processes may take place at different scales at the same time, and that the scales of a study must be chosen in accordance with the objectives pursued. Apart from conventional approaches to genetic structure, spatial autocorrelation and related techniques, such as Mantel test, correlograms, Mantel correlograms, join-counts, variograms and point pattern analysis, can detect and characterise the existence of spatial genetic structures and lead the way to discussing the environmental and biological factors responsible for them. An alternative way of including spatial variability in modelling approaches that deal with genetic patterns or processes is through the use of constrained ordinations. Although scarcely used at present, these methodologies have great applicability in conservation biology and can lead a way to an effective integration of genetic, demographic and ecological perspectives.

Shrub encroachment can reverse desertification in semi-arid Mediterranean grasslands

The worldwide phenomenon of shrub encroachment in grass-dominated dryland ecosystems is commonly associated with desertification. Studies of the purported desertification effects associated with shrub encroachment are often restricted to relatively few study areas, and document a narrow range of possible impacts upon biota and ecosystem processes. We conducted a study in degraded Mediterranean grasslands dominated by Stipa tenacissima to simultaneously evaluate the effects of shrub encroachment on the structure and composition of multiple biotic community components, and on various indicators of ecosystem function. Shrub encroachment enhanced vascular plant richness, biomass of fungi, actinomycetes and other bacteria, and was linked with greater soil fertility and N mineralization rates. While shrub encroachment may be a widespread phenomenon in drylands, an interpretation that this is an expression of desertification is not universal. Our results suggest that shrub establishment may be an important step in the reversal of desertification processes in the Mediterranean region.

Structural equation modelling: an alternative for assessing causal relationships in threatened plant populations

Structural equation modelling (SEM) is a powerful tool to explore and contrast hypotheses on causal relationships among variables using observational data. It constitutes an alternative to experimental approaches that is especially useful in the conservation of small populations where the implementation of treatments may have a negative effect on population viability. We are presently applying SEM to study the factors that condition reproductive success, seed emergence and plantlet survival in several plant species. We are also using model comparisons through multi-sample analysis to assess the implications of different microhabitats on the viability of a population. The most outstanding advantages of this tool are the global perspective used in the study of complex problems, the ability to discern the essential from the accessory, and the possibility of evaluating ones own hypotheses. The basic procedure, the limitations of this method and further applications in conservation and management are also discussed.

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.

Factors affecting establishment of a gypsophyte: the case of Lepidium subulatum (Brassicaceae)

The restriction of vascular plants to gypsum-rich soils under arid or semiarid climates has been reported by many authors in different parts of the world. However, factors controlling the presence of gypsophytes on these soils are far from understood. We investigated the establishment of Lepidium subulatum, a gypsophyte, in a nondisturbed semiarid gypsum-soil landscape in central Spain, both from spatial and temporal perspectives. Over 1400 seedlings were tagged, and their growth and survival were monitored for a 2-yr period. Several biotic and abiotic variables were measured to determine the factors controlling the emergence and early survival. These variables included the cover of annual plants, bryophytes, lichens, litter, gypsum crystals, bare fraction and cover of each perennial plant, and several soil properties (gravel, fine gravel, and fine-earth fraction, conductivity, pH, gypsum content, organic matter and penetrometer soil resistance). Our results support the linkage of gypsophily with some physical properties of the surface crust. Seedlings tended to establish on the gypsum surface crust, and their survival was size dependent, probably as a consequence of the necessity of rooting below the surface crust before summer drought arrives. However, once seedlings emerged, a higher survival rate occurred on the alluvial soils of the piedmont-slope boundary where soil crusts are absent or thinner. We conclude that Lepidium subulatum may be considered a refuge model endemic with a distribution range that occupies a reduced fraction of a wider habitat from which it is probably excluded by competition.

Inhibitory effects of Artemisia herba-alba on the germination of the gypsophyte Helianthemum squamatum

The potential allelopathic role of Artemisia herba-alba has been evaluated in order to explain the community pattern of the gypsum semiarid environments of central Spain. This pattern shows a sharp ecotone between a gypsophile sparse shrubby community dominated by Helianthemum squamatum, which grows on slopes with gypsum surface crusts, and a nitrohalophilous community on the gypsum alluvial soils of piedmont dominated by Artemisia herba-alba. In order to explain this pattern, resource limitation was discarded because no significant differences in several soil parametrers, but fine earth fraction and organic matter content, had been detected in a previous study. Results confirm the inhibitory effect of aqueous extracts on the final germination percentage of scarified seeds of Helianthemum squamatum and also on the shape of the germination curves, which indicate delay of germination for the aqueous extracts. Assays with soil of the Artemisiacommunity (three types) showed that germination was strongly inhibited in soils obtained below the canopy of mature plants (litter maintained) and retarded in the other treatments. We suggest that the spatial community pattern detected in this gypsum environment and characterised by a sharp ecotone could be at least partially controlled by interference through allelopathy. This determines the excliusion of Helianthemum squamatum plants from alluvial soils.

Small-scale patterns of abundance of mosses and lichens forming biological soil crusts in two semi-arid gypsum environments

Despite important advances in the understanding of biological soil crusts and their key role in ecosystem processes in arid and semi-arid environments, little is known about those factors driving the small-scale patterns of abundance and distribution of crust-forming lichens and mosses. We used constrained ordination techniques (RDAs) to test the hypothesis that the spatial patterning of lichens and mosses is related to surface and subsurface soil variables in two semi-arid gypsum environments of Spain. Our results show that the abundance of mosses and lichens forming biological soil crusts was related to a limited set of variables (cover of bare soil and litter, soil respiration, potassium content and aggregate stability). Moreover, they provide some insights into the importance of these variables as drivers of biological soil-crust composition and abundance in semi-arid gypsum environments.

Seed bank spatial pattern in a temperate secondary forest

Abstract Seed bank spatial pattern was studied in a secondary forest dominated by Fagus sylvatica and Betula celtiberica in the Urkiola Natural Park (N Spain). Soil samples were taken every 2 m in a regular grid (196 points) and divided into two fractions (0-3 cm and 3-10 cm deep). The viable seed bank was studied by monitoring seedling emergence for ten months. The effect of different factors on seed bank composition and patterning was analysed using constrained ordination as a hypothesis testing tool. Furthermore, the existence of spatial autocorrelation was evaluated by geostatistical analysis. Seed density was high, 7057 seed.m−2, with a few species dominating. Species composition in the various layers were significantly correlated. The seed bank showed significant spatial structure, which was partially explainable by the spatial structure of the canopy and understorey vegetation. Spatial clumping from 0-8 m was observed in seed bank density and composition, mainly due to the pattern of two abundant taxa Juncus effusus and Ericaceae. The Ericaceae seed bank was related to the spatial distribution of dead stumps of Erica arborea. J. effusus was not present in the above-ground vegetation, which indicates that its seed bank was formed in the past. As expected, the seed bank of this forest reflects its history, which is characterized by complex man-induced perturbations. The seed bank appears to be structured as a consequence of contrasting driving forces such as canopy structure, understorey composition and structural and microhabitat features. Abbreviations: TVE = Total variance explained. Nomenclature: Aizpuru et al. (1999).

Edge effects on epiphytic communities in a Mediterranean Quercus pyrenaica forest

Abstract Question: What are the edge effect responses of epiphytic lichen communities in Mediterranean Quercus pyrenaica forest? Location: Central Spain. Methods: We established ten transects perpendicular to a road dissecting a well conserved remnant of Q. pyrenaica forest into two sections. Transects extended from the forest/road edge to 100 m into the forest. Data were collected from seven plots in each transect at different distances from the edge. Variables were grouped into stand scale variables (distance to edge, number of trees per plot, mean diameter per plot, irradiance) and tree scale variables (diameter and height of sampled trees, aspect of the sampled square and relative height of the square). We used General Mixed Linear Models and constrained ordination techniques to test the hypothesis that the spatio-temporal heterogeneity of light and water controls the occurrence of lichens and bryophytes along the edge-interior gradient in the Q. pyrenaica forest. Results: Microclimatic parameters vary in a non-linear way; edge and interior stands showed the most divergent and extreme values. Although the micro-environment within Mediterranean forests is heterogeneous, interior conditions are apparently suitable for the performance of some specific forest epiphytes. Consequently, species richness does not show significant differences along the gradient. Total epiphytic cover increases towards the forest interior, but distance to the edge together with other predictors at the tree scale (aspect and height of the square) are the most relevant predictors for the composition and structure of these communities. Conclusions: Composition and structure of epiphytic communities in a Mediterranean semi-deciduous forest are affected by the edge between the forest and the road constructed. Since some extremely rare lichens only occur at interior stands, the conservation of these threatened elements requires urgent conservation measures because well preserved and unmanaged forests in the Mediterranean region are very rare. Nomenclature: Hafellner & Türk (2001); Bisby & Rostov (2005).