Yolanda Pueyo

Spatial vegetation patterns and imminent desertification in Mediterranean arid ecosystems

Humans and climate affect ecosystems and their services, which may involve continuous and discontinuous transitions from one stable state to another. Discontinuous transitions are abrupt, irreversible and among the most catastrophic changes of ecosystems identified. For terrestrial ecosystems, it has been hypothesized that vegetation patchiness could be used as a signature of imminent transitions. Here, we analyse how vegetation patchiness changes in arid ecosystems with different grazing pressures, using both field data and a modelling approach. In the modelling approach, we extrapolated our analysis to even higher grazing pressures to investigate the vegetation patchiness when desertification is imminent. In three arid Mediterranean ecosystems in Spain, Greece and Morocco, we found that the patch-size distribution of the vegetation follows a power law. Using a stochastic cellular automaton model, we show that local positive interactions among plants can explain such power-law distributions. Furthermore, with increasing grazing pressure, the field data revealed consistent deviations from power laws. Increased grazing pressure leads to similar deviations in the model. When grazing was further increased in the model, we found that these deviations always and only occurred close to transition to desert, independent of the type of transition, and regardless of the vegetation cover. Therefore, we propose that patch-size distributions may be a warning signal for the onset of desertification.

Quantitative characterization of the regressive ecological succession by fractal analysis of plant spatial patterns

We studied the effect of grazing on the degree of regression of successional vegetation dynamic in a semi-arid Mediterranean matorral. We quantified the spatial distribution patterns of the vegetation by fractal analyses, using the fractal information dimension and spatial autocorrelation measured by detrended fluctuation analyses (DFA). It is the first time that fractal analysis of plant spatial patterns has been used to characterize the regressive ecological succession. Plant spatial patterns were compared over a long-term grazing gradient (low, medium and heavy grazing pressure) and on ungrazed sites for two different plant communities: A middle dense matorral of Chamaerops and Periploca at Sabinar-Romeral and a middle dense matorral of Chamaerops, Rhamnus and Ulex at Requena-Montano. The two communities differed also in the microclimatic characteristics (sea oriented at the Sabinar-Romeral site and inland oriented at the Requena-Montano site). The information fractal dimension increased as we moved from a middle dense matorral to discontinuous and scattered matorral and, finally to the late regressive succession, at Stipa steppe stage. At this stage a drastic change in the fractal dimension revealed a change in the vegetation structure, accurately indicating end successional vegetation stages. Long-term correlation analysis (DFA) revealed that an increase in grazing pressure leads to unpredictability (randomness) in species distributions, a reduction in diversity, and an increase in cover of the regressive successional species, e.g. Stipa tenacissima L. These comparisons provide a quantitative characterization of the successional dynamic of plant spatial patterns in response to grazing perturbation gradient.

Gypsophile vegetation patterns under a range of soil properties induced by topographical position

Iberian gypsophile plant communities are considered a priority for conservation by the European Community because of their highly specialized flora in gypsum outcrops in arid and semiarid regions. Despite the ecological importance of these ecosystems, the edaphic factors that constrain plant communities on gypsiferous soils remain unclear. It has been proposed that both the chemical and physical restrictive conditions of gypsum soils determine gypsophily in plants. Here we hypothesize that the rigors of the gypsum soil environment depends on topography, decreasing from flat areas on hilltops to south-oriented slopes and finally to slopes oriented to the north. We also hypothesized that the relaxation of the rigors of the gypsum soil environment with topography affects both to individual plant and community characteristics of gypsophile vegetation: we expect a reduction of gypsophyte abundance, an increase of diversity and the amelioration of facilitative interactions of plant species. We analysed the physical and chemical properties of gypsum soils that have been proposed that determine the rigors of the gypsum soil environment (i.e.: unbalanced ion concentrations and superficial soil crust). The predicted rigor gradient along topographical locations was confirmed and was mainly caused by superficial soil crust. The decreasing rigor gradient was accompanied by a fall in the abundance of gypsophytes. However, when gypsophytes were considered separately, several patterns were observed, indicating distinct tolerance to relaxation of rigor of the gypsum soil conditions and different competition abilities between gypsophytes. Plant species were more clumped, and gypsophile communities presented higher diversity, evenness and richness values where rigor of gypsum soil conditions were maximum (flat hilltop positions). Relaxation of rigor (north-oriented slopes) was characterized by loss of facilitative interaction between species and the dominance of the gypsovag Rosmarinus officinalis L., although richness was still very high, which can be attributed to the coexistence of gypsophytes and gypsovags. We conclude that the rigor of gypsum soil environment gradient with topography is mainly determined by superficial soil crust, and it is a crucial determinant of gypsophile plant communities.

Is the patch size distribution of vegetation a suitable indicator of desertification processes? Comment.

With ongoing climate change, the search for indicators of imminent ecosystem shifts is attracting increasing attention (e.g., Scheffer et al. 2009). Recently, the spatial organization of ecosystems has been suggested as a good candidate for such an indicator in spatially structured ecosystems (Rietkerk et al. 2004, Ke´fi et al. 2007a, Guttal and Jayaprakash 2009).

DIFFERENCES IN GYPSUM PLANT COMMUNITIES ASSOCIATED WITH HABITAT FRAGMENTATION AND LIVESTOCK GRAZING

The negative consequences of habitat fragmentation for plant communities have been documented in many regions of the world. In some fragmented habitats, livestock grazing has been proposed to be a dispersal mechanism reducing isolation between fragments. In others, grazing acts together with fragmentation in a way that increases habitat degradation. Iberian gypsum plant communities have been grazed and fragmented by agricultural practices for centuries. Although their conservation is considered a priority by the European Community, the effects of fragmentation on gypsum plant communities and the possible role of livestock grazing remain unknown. In addition, a substantial proportion of plant species growing in gypsum environments are gypsum specialists. They could be particularly affected by fragmentation, as was found for other habitat specialists (i.e., serpentine and calcareous specialists). In this study (1) we investigated the effect of fragmentation and grazing on gypsum plant community composition (species and life-forms), and (2) we tested to see if gypsum specialists were differently affected by fragmentation and grazing than habitat generalists. A vegetation survey was conducted in the largest gypsum outcrop of Europe (Middle Ebro Valley, northeast Spain). Fragmented and continuous sites in grazed and ungrazed areas were compared. Measurements related to species and composition of life-forms were contrasted first for the whole gypsum plant community and then specifically for the gypsum specialists. In the whole community, our results showed lower plant species diversity in fragmented sites, mainly due to the larger dominance of species more tolerant to fragmented habitat conditions. With livestock grazing, the plant species richness and the similarity in plant species composition between remnants was larger, suggesting that animals were acting as dispersal agents between fragments. As expected, gypsum specialists were less abundant in fragmented areas, and grazing led to the disappearance of the rare gypsum specialist Campanula fastigiata. According to our results, conservation strategies for gypsum plant communities in human-dominated landscapes should consider that fragmentation and grazing modify plant community composition affecting gypsum specialists in particular.

Fractal analysis of plant spatial patterns: a monitoring tool for vegetation transition shifts

Abstract.Spatial heterogeneity, like species diversity, is an important ecosystem property. We examine the effects of land use on the diversity and spatial distribution of plants in five semi-arid communities of eastern Spain using non-linear methods to assess the spatialtemporal dynamics of plant populations. Specifically, we are interested in detecting long-term structural changes or drift in an ecosystem before it is too late to prevent irreversible degradation. Fractal analysis is used to characterize the complexity of plant spatial patterns and Information Theory indices are used to measure change in information flow with land use changes and soil substrate. We found that grazing favored diversity and heterogeneity of species distribution on the impoverished gypsum and saline substrate community, as opposed to the detrimental effect of grazing in the Alpha steppe community. Indeed, old-field succession after 30 years of abandonment showed a recovery of species diversity but not the spatial structure of the vegetation. Further, Information Fractal Dimension, representing the unpredictability of plant spatial patterns in the landscape, increased as we moved from a highly diverse to a less diverse community, revealing the change to a more scattered and homogeneous spatial plant distribution. The Information Fractal Dimension is a good estimator of ecosystem disturbance, independent of scale, and thus can be used to monitor ecosystem dynamics.

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.

The role of reproductive plant traits and biotic interactions in the dynamics of semi-arid plant communities.

The dynamics of semi-arid plant communities are determined by the interplay between competition and facilitation among plants. The sign and strength of these biotic interactions depend on plant traits. However, the relationships between plant traits and biotic interactions, and the consequences for plant communities are still poorly understood. Our objective here was to investigate, with a modelling approach, the role of plant reproductive traits on biotic interactions, and the consequences for processes such as plant succession and invasion. The dynamics of two plant types were modelled with a spatially-explicit integrodifferential model: (1) a plant with seed dispersal (colonizer of bare soil) and (2) a plant with local vegetative propagation (local competitor). Both plant types were involved in facilitation due to a local positive feedback between vegetation biomass and soil water availability, which promoted establishment and growth. Plants in the system also competed for limited water. The efficiency in water acquisition (dependent on reproductive and growth plant traits) determined which plant type dominated the community at the steady state. Facilitative interactions between plant types also played an important role in the community dynamics, promoting establishment in the driest conditions and recovery from low biomass. Plants with vegetative propagation took advantage of the ability of seed dispersers to establish on bare soil from a low initial biomass. Seed dispersers were good invaders, maintained high biomass at intermediate and high rainfall and showed a high ability in taking profit from the positive feedback originated by plants with vegetative propagation under the driest conditions. However, seed dispersers lost competitiveness with an increasing investment in fecundity. All together, our results showed that reproductive plant traits can affect the balance between facilitative and competitive interactions. Understanding this effect of plant traits on biotic interactions provides insights in processes such as plant succession and shrub encroachment.

Determinants of land degradation and fragmentation in semiarid vegetation at landscape scale

The objective of this paper was to investigate the sensitivity to degradation of semiarid plant communities in terms of plant cover and fragmentation, quantifying relationships between landscape characteristic (physical, socio-economical and historical) and vegetation degradation. The degradation of vegetation was measured as the degree of coverage of the two dominant vegetation types (i.e. tall arid brush and tall grass steppe), while fragmentation was measured as patch size and isolation. Data were obtained using GIS tools, and analyzed by logistic regression and linear multivariate regression. Results showed denser coverage at more elevated, gradual slopes that were not sea-oriented. Historical elements of the landscape had a significant effect on current natural vegetation. According to the fragmentation patterns, the vegetation is in fairly good condition (medium coverage had the largest patches but dense coverage was less isolated) but attention must be given to preserve vegetation, due to the relationships between fragmentation and human activities. Moreover, the protection plan under way in the area appeared to favour denser vegetation cover, while human activities had a measurable effect on vegetation degradation.

Plant–plant interactions as a mechanism structuring plant diversity in a Mediterranean semi‐arid ecosystem

Abstract Plant–plant interactions are among the fundamental processes that shape structure and functioning of arid and semi‐arid plant communities. Despite the large amount of studies that have assessed the relationship between plant–plant interactions (i.e., facilitation and competition) and diversity, often researchers forget a third kind of interaction, known as allelopathy. We examined the effect of plant–plant interactions of three dominant species: the perennial grass Lygeum spartum, the allelopathic dwarf shrub Artemisia herba‐alba, and the nurse shrub Salsola vermiculata, on plant diversity and species composition in a semi‐arid ecosystem in NE Spain. Specifically, we quantified the interaction outcome (IO) based on species co‐occurrence, we analyzed diversity by calculation of the individual species–area relationship (ISAR), and compositional changes by calculation of the Chao‐Jaccard similarity index. We found that S. vermiculata had more positive IO values than L. spartum, and A. herba‐alba had values between them. Lygeum spartum and A. herba‐alba acted as diversity repellers, whereas S. vermiculata acted as a diversity accumulator. As aridity increased, A. herba‐alba transitioned from diversity repeller to neutral and S. vermiculata transitioned from neutral to diversity accumulator, while L. spartum remained as diversity repeller. Artemisia herba‐alba had more perennial grass species in its local neighborhood than expected by the null model, suggesting some tolerance of this group to its “chemical neighbor”. Consequently, species that coexist with A. herba‐alba were very similar among different A. herba‐alba individuals. Our findings highlight the role of the nurse shrub S. vermiculata as ecosystem engineer, creating and maintaining patches of diversity, as well as the complex mechanism that an allelopathic plant may have on diversity and species assemblage. Further research is needed to determine the relative importance of allelopathy and competition in the overall interference of allelopathic plants.