Mohamed Chaieb

Decoupling of soil nutrient cycles as a function of aridity in global drylands

The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability. The increase in aridity predicted for the twenty-first century in many drylands worldwide may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

Life history strategies and water resource allocation in five pasture species of the Tunisian arid zone

Abstract Five sympatric species of perennial pasture plants were studied in monospecific plots at an experimental site in the arid zone of southern Tunisia. For one year following establishment, irrigation was supplied monthly to test plots so as to complement natural precipitation and simulate an “optimum”; rain year (250 mm of precipitation occurring, evenly distributed, between September and April). In addition to hydrometric fluctuations in the upper soil layers, growth rate and phenological activity were monitored for all species (Cenchrus ciliaris, Digitaria commutata, Stipa lagascae (Poaceae), Argyrolobium uniflorum (Fabaceae) and Plantago albicans (Plantaginaceae)). Overall growth rate and foliar extension measurements revealed differential water use efficiency, phenology, and response to water stress in the species studied. C. ciliaris and D. commutata were the most efficient in their use of water. The former, a C4 grass species of paleo‐tropical origin, was primarily active in spring and autumn....

Climate and soil attributes determine plant species turnover in global drylands

AIM Geographic, climatic, and soil factors are major drivers of plant beta diversity, but their importance for dryland plant communities is poorly known. This study aims to: i) characterize patterns of beta diversity in global drylands, ii) detect common environmental drivers of beta diversity, and iii) test for thresholds in environmental conditions driving potential shifts in plant species composition. LOCATION 224 sites in diverse dryland plant communities from 22 geographical regions in six continents. METHODS Beta diversity was quantified with four complementary measures: the percentage of singletons (species occurring at only one site), Whittakes beta diversity (β(W)), a directional beta diversity metric based on the correlation in species occurrences among spatially contiguous sites (β(R2)), and a multivariate abundance-based metric (β(MV)). We used linear modelling to quantify the relationships between these metrics of beta diversity and geographic, climatic, and soil variables. RESULTS Soil fertility and variability in temperature and rainfall, and to a lesser extent latitude, were the most important environmental predictors of beta diversity. Metrics related to species identity (percentage of singletons and β(W)) were most sensitive to soil fertility, whereas those metrics related to environmental gradients and abundance ((β(R2)) and β(MV)) were more associated with climate variability. Interactions among soil variables, climatic factors, and plant cover were not important determinants of beta diversity. Sites receiving less than 178 mm of annual rainfall differed sharply in species composition from more mesic sites (> 200 mm). MAIN CONCLUSIONS Soil fertility and variability in temperature and rainfall are the most important environmental predictors of variation in plant beta diversity in global drylands. Our results suggest that those sites annually receiving ~ 178 mm of rainfall will be especially sensitive to future climate changes. These findings may help to define appropriate conservation strategies for mitigating effects of climate change on dryland vegetation.

Acacia saligna Plantation Impact on Soil Surface Properties and Vascular Plant Species Composition in Central Tunisia

Exotic species have been widely used in reforestation in arid areas in northern Africa, as they provide ecosystem services that native species may be unable to supply. But information on their effect on ecosystem function and composition is scarce. We have evaluated the effect of 20-year-old Acacia saligna (Labill.) H.L. Wendl. plantations on the soil surface conditions and vascular plant composition in central Tunisia. Landscape function analysis (LFA) indices of soil stability, infiltration, and nutrient recycling underneath A. saligna increased by 1%, 138%, and 148%, respectively, in comparison to bare areas. This species, however, did not affect indicators of soil surface functionality, the spatial pattern of resource sinks-sources, or vascular plant species composition at a plot scale. This was probably due to low A. saligna cover, and a limited degree of litter incorporation into the soil surface. Our results suggest that the potential for improving ecosystem function and fostering succession by using this species seems to be limited in this arid environment.

The relative contribution of short-term versus long-term effects in shrub-understory species interactions under arid conditions

Abstract Plant–plant interactions (competition and facilitation) in terrestrial ecosystems include: (1) short-term effects, primarily quantified with experimental removals; and (2) long-term effects, mostly due to soil weathering processes, primarily quantified with observational methods. It has been argued that these effects are likely to vary in contrasting directions with increasing drought stress in arid systems. However, few studies have used appropriate methodology to assess both types of effects and their variation across nurse species and environmental conditions, in particular in arid systems. This knowledge is crucial for predicting variation in the mediating role of facilitation with climate change and assessing the importance of nurse effects in ecological restoration. In the arid climate of central-south Tunisia, understory species’ biomass, abundance and composition and soil parameters were compared in shrub-control, shrub-removed and open areas for three shrub species and in two habitats with contrasting soil moisture conditions. Long-term effects were dominant, positive for all three shrub species and associated with increasing nutrient content in shrub patches. Short-term effects, mainly related to water consumption, were weaker, mostly negative and dependent on shrub species. Additionally, long-term effects were less positive and short-term effects more negative in the dry habitat than in the wet habitat. Our study provides evidence of the primary influence of positive (facilitative) long-term effects in this arid system. However, the net effects of shrubs could be less beneficial for other species with increasing aridity under climate change, due to both a decrease in positive long-term effects and an increase in negative short-term effects.

Screening biological traits and fluoride contents of native vegetations in arid environments to select efficiently fluoride-tolerant native plant species for in-situ phytoremediation

High fluoride pollution has been detected in the surrounding soils of the coastal superphosphate industries in the Gulf of Gabes (Southeast of Tunisia). A study was conducted in vicinity of factories analysing plant functional traits combined with plant fluoride accumulation and soil metal concentrations aiming to screen more efficiently native plant species tolerant to this pollution. Aerial parts of 18 plant species out of the 10 most abundant species per site were harvested on two polluted sites of Gabes and Skhira at the vicinity of the factories and on the less polluted site of Smara. Native plant species accumulated fluoride following the gradient of soil pollution. Fluoride contents of plant aerial parts ranged from 37 mg kg(-1) to 360 mg kg(-1) and five plant species were only found in the most polluted site. However these latter had low biomass and soil cover. Crossing biological traits and fluoride contents, a selection grid for potentially restorative plant species enabled the selection of three native perennials i.e. Rhanterium suaveolens, Atractylis serratuloides and, Erodium glaucophyllum as potential candidates for an in-situ phytoremediation program on arid fluoride-polluted sites. This approach may be used in other fluoride-polluted Mediterranean environments.

Aboveground phytomass productivity and morphological variability of Tunisian accessions of Cenchrus ciliaris L.

This study assessed the aboveground aerial phytomass produced by 47 accessions of Cenchrus ciliaris L. locally collected in the arid zone of Tunisia and classified into six eco-geographic groups. The extent of genetic variation in five morphologic parameters was examined. Correlations among all parameters were also analysed. For three years, phytomass of these accessions was cut and measured for three cutting periods characterised by different rainfall features. In 1998 the mean of the phytomass productivity was 199.1g DM (Dry Matter) plant–1. The productivity in 1999 was 81.0g DM plant–1, but in 2000 the productivity was only 21.6g DM plant–1. Positive correlations were found between dry matter and all morphological parameters, especially tuft diameter (r = 0.670) and tuft height (r = 0.637). Principal components analysis was carried out on all variables. For both dry matter and phenology, individuals differentiated strongly, but irrespective of their accession or eco-geographic origin.

Effects of water stress and soil type on photosynthesis, leaf water potential and yield of olive trees (Olea europaea L. cv. Chemlali Sfax)

In Tunisia, olives are grown under severe rain-fed, arid conditions. To determine the behaviour of olive trees (cv. Chemlali Sfax) during the severe drought affecting Tunisian arid areas in 2002, a range of physiological parameters were investigated in three adjacent orchards. Two olive orchards were rain-fed, one located on a sandy soil, and the other on a sandy-loam clay soil. A third orchard was also located on sandy soil, but received remedial irrigation (415 mm of water per year; ~40% of olive evapotranspiration). Predawn leaf water potential (Ψpd) did not fall below –1.52 MPa for irrigated olive trees. However, a large decrease in Ψpd was observed for rain-fed olive trees in the same period with Ψpd measured at about –3.2 MPa on sandy soil and –3.6 MPa on sandy-loam clay soil. At the same time, the minimal leaf water potential recorded at midday (Ψmin) decreased to –4.15 MPa and –4.71 MPa in the rain-fed trees for sandy and sandy-loam clay soil, respectively. For irrigated trees, the Ψmin was –1.95 MPa. These results were associated with relative water content, which varied from 80% for irrigated trees to 54 and 43.6%, respectively, for rain-fed trees and trees subjected to severe drought. In August, when the relative water content values were less than 50%, a progressive desiccation in the outer layer of canopy and death of terminal shoots were observed in trees, which grew on the sandy-loam clay soil. Furthermore, low soil water availability also affected (negatively) the net photosynthetic rate in rain-fed orchards (10.3 µmol/m2.s for irrigated trees v. 5.3 µmol/m2.s in rain-fed trees on sandy soil) and stomatal conductance (98.5 mmol/m2.s v. 69.3 mmol/m2.s). However, it improved water use efficiency (7.6 v. 4.7 µmol CO2/mmol H2O), which increased by more than 50% in both groups of rain-fed trees compared with the irrigated ones. We can conclude that olive trees respond to drought by showing significant changes in their physiological and biological mechanisms. These results also help our understanding of how olive trees cope with water stress in the field and how marginal soils can restrict growth and lower yields.

New insights into the polyploid complex Cenchrus ciliaris L. (Poaceae) show its capacity for gene flow and recombination processes despite its apomictic nature

Cenchrus ciliaris L. is a C4 perennial grass of arid lands which is under the focus of different ecological issues such as response to desertification, quality of forage grass and impacts of invasions. Here, molecular and morphological analyses of the genetic diversity of several Tunisian provenances of C. ciliaris were performed to better understand the phenotypic polymorphism of this agamospermous and polyploid grass. Ten phenotypic traits associated with productivity were measured in a common garden environment. Amplified Fragment Length Polymorphism (AFLP) markers were developed to investigate the structure of genetic diversity among and within provenances and between the three ploidy levels. Heritable phenotypic traits showed considerable differences within provenances. Surprisingly, AFLP markers revealed the existence of genotypic variations between individuals of the same sibship and a high G/N value (0.55). A neighbour-joining tree based on AFLP markers revealed three major groups; tetraploid, pentaploid and a mix of pentaploid and hexaploids. These groups do not correspond completely to the geographical origin of samples. The results underline the possibility of sexual reproduction, recombination and gene flow within and between populations of C. ciliaris. In respect with the well known dynamic nature of polyploid genomes, these results should have strong consequences for the future management of this grass for both conservation and invasion issues.

Using trees as a restoration tool in Tunisian arid areas: effects on understorey vegetation and soil nutrients

A field experiment was conducted in an afforested Stipa tenacissima L. steppe in arid southern Tunisia to evaluate the effect of three tree species (Acacia salicina Lindl., Pinus halepensis Mill. and Eucalyptus occidentalis Endl.) on understorey vegetation and soil nutrients. For each tree species, two subhabitats were distinguished: under the canopy, and out in the open. Organic carbon, total N, available P and pH were higher under the canopies of the three tree species than out in the open, and the effect was more pronounced in the top 10 cm of soil. Similarly, plant cover, biomass, richness and diversity were significantly higher under tree canopies. Some species such as Plantago amplexicaulis Cav., Helianthemum kahiricum Del. and Artemisia campestris L., which use large amounts of soil nutrients, showed a strong preference for areas under the canopy. Among the three tree species, Acacia salicina had the strongest positive effect on soil nutrients and understorey vegetation, and, thus, may be more useful for restoring arid areas and creating areas of enhanced soil nutrients than Pinus or Eucalyptus.