Competition for microsites during recruitment in semiarid annual plant communities.

The concept of microsites for recruitment is central to plant ecology, but it is unclear whether these sites are abstract constructs or real entities. I hypothesize that, in generally microsite-limited communities, microsites comprise a limiting physical resource for which different species compete. I tested this hypothesis on winter-annual communities on biocrust in the semiarid Northern Negev of Israel, in which most species are microsite-limited, while the dominant grass (Stipa capensis) has overcome this limitation by efficient microsite acquisition and a lack of secondary seed dormancy. I tested whether the dominant suppresses the subordinate species, collectively, during recruitment, rather than during growth. To this end, I conducted a field experiment with three blocks of six plots (6 m × 6 m) with two treatments - mowing in spring 2006 (intershrub, intershrub + shrub patches, and none) and shrub-patch removal (0% or 50% of the patches). I collected data from four seed traps per plot before spring 2007 and from five plant samples per plot at the end of spring. Mowing significantly reduced both seed and plant density of the dominant species, reflecting seed-limited recruitment, and increased subordinate plant density by competitive release. Multiple regressions of per-plant and per-gram effects and responses showed that competition was a direct effect of the dominant's density. Total and per-group biomass was proportional to density, implying density-independent per capita growth. Subordinate species number also increased with their density, due to the sample-size effect. These findings indicate that the seed-limited dominant diffusely suppresses the subordinates during recruitment, supporting the microsite competition hypothesis. The shift from growth resources to microsites extends the role of inter-specific competition along productivity and disturbance gradients, and highlights the asymmetric relationship between the two kinds of competition, as microsite competition is only observable if initial abundances are not overshadowed by density-dependent growth and mortality. The findings also demonstrate that (1) lacking secondary seed dormancy is an evolutionarily stable strategy in dryland annuals, alongside seed dormancy in microsite-limited species, and (2) biomass removal (e.g., by herbivory) increases small-scale biodiversity, enhancing the sustainability of dryland grazing, but without compensatory growth.

Climate change scenarios of herbaceous production along an aridity gradient: vulnerability increases with aridity.

Climate change is expected to reduce annual precipitation by 20% and increase its standard deviation by 20% in the eastern Mediterranean. We have examined how these changes may affect herbaceous aboveground net primary production (ANPP) and its inter-annual coefficient of variation (CV) in natural rangelands along a desert-Mediterranean precipitation gradient, at five sites representing arid, semi-arid, and Mediterranean-type ecosystems, respectively, all showing positive linear relationships between herbaceous ANPP and annual precipitation. Scenarios of reduced annual precipitation and increased inter-annual precipitation variability were defined by manipulating mean annual precipitation (MAP) and its standard deviation. We simulated precipitation and calculated ANPP using current ANPP-precipitation relationships. Our model predicts that reduced precipitation will strongly reduce ANPP in arid and semi-arid sites. Moreover, the effect of reduced precipitation on the CV of ANPP along the entire gradient may be modified by changes in inter-annual variability in MAP. Reduced precipitation combined with increased precipitation variability was the scenario most relevant to the wet end of the gradient, due to the increased likelihood for both dry and rainy years. In contrast, the scenario most relevant to the arid end of the gradient combined reduced precipitation with decreased precipitation variability, due to the strong effect on mean ANPP. All scenarios increased variability of ANPP along the entire gradient. However, the higher sensitivity of vegetation at arid and semi-arid sites (i.e., lower forage production) to future changes in the precipitation regime emphasizes the need to adapt grazing management in these ecosystems to secure their long-term viability as sustainable rangelands.

Plant species richness and ecosystem multifunctionality in global drylands.

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.

Annual plant community responses to density of small-scale soil disturbances in the Negev Desert of Israel.

We investigated whether plant diversity and productivity in small-scale soil disturbances, which is known to be higher than in undisturbed soil, decreases as the density of the disturbances increases. We studied this in an experiment with soil diggings (15 cm diameter and 15 cm depth) dug at a range of densities, on a north- and a south-facing slope of a watershed in the central Negev Desert of Israel. The diggings were similar to the commonly occurring pits made by porcupines (Hystrix indica) as they forage for below-ground plant parts. We used four levels of digging density, within the naturally occurring range in the region, represented by a rectangular plot with rows of diggings dug at four distances between diggings. The plots were laid out in a blocked design with three replications on both slopes, with each block containing all four levels of digging density. In the spring of 1992, 1994 and 1995 we measured plant density, species richness and plant productivity in the diggings, and in adjacent equal-sized undisturbed control areas ("soil matrix") and on the mounds made by the removed excess soil. Plant density, species richness and productivity of annual plants were higher in the diggings than in the undisturbed matrix, while these responses were very low on the mounds. Plant density, species richness and productivity in the diggings, but not in the matrix or mounds, decreased as digging density increased. This effect varied slightly with location within a watershed and with annual rainfall. The density of seeds captured in the diggings from outside the digging during the 1995 dispersal season decreased with increasing digging density, but only on one of the slopes. At the highest digging density, plant density and species number in the diggings did not decrease down the slope, as expected if interference between diggings in runoff water capture were the cause of the digging density effect. There was a weak decrease in biomass production in 1994-1995 down the slope. We used a simple mathematical model to estimate whether the distribution of rainfall intensities that occurred during the winter of 1994-1995 could result in differences between digging densities in the amount of water captured by the diggings, and whether this could explain the observed effect of digging density. The model showed that there were four events during which less water was captured by the diggings at high digging densities, except in the topmost row of diggings. Soil moisture measurements, however, showed very little difference between diggings at different digging densities. We explain our findings as the result of the interaction between the properties of the disturbance patch with its surroundings, as the diggings capture resources in the form of runoff water, and seeds moved primarily by wind. The additional resources and seeds captured in diggings increase plant density, species richness and productivity relative to the undisturbed matrix. However, the contrast in plant responses between the disturbed patches and undisturbed soil diminishes at higher digging densities. We explain this as interference among diggings at close proximity. As we did not detect a decrease in plant responses down the slopes, we conclude that interference is due to interception of the wind-driven, non-directional flow of seeds. Interception of the down-slope flow of runoff water by upslope diggings is insufficient to affect plant density, determined at the beginning of the season. Later in the season, runoff interception may become important for biomass production.

Life histories of two desert species of the bulbous genus Bellevalia : The relation between biomass partitioning and water availability.

Annual biomass increment and biomass partitioning to leaves, roots and reproduction, and biomass storage in the below-ground bulb was measured in plants of two species of the geophytic genus Bellevalia grown outdoors at three levels of soil moisture. The differences between the species were in accordance with the hypothesis that plants of more arid environments should rely more on internal reserves than plants of more productive environments. In Bellevalia desertorum, a shallow rooted species of the most arid habitats in the Central Negev, leaf and root development during outgrowth at the beginning of winter was rather variable, and followed soil moisture availability to a certain degree. A small portion of its biomass budget was committed to seed production, which varied little among the irrigation regimes. The rest of the biomass was stored in the bulb. The amount of biomass devoted annually to reproduction was mainly determined by the amount of reserves already present in the bulb. In contrast, in B. eigii, which grows in the more productive wadis with its bulb at a depth of 15 to 30 cm, leaf and root growth was not only determined by water availability, but also by initial bulb mass. This resulted in a greater potential relative growth rate than in B. desertorum, but also in a greater risk of accumulating less biomass than it spent in root and leaf construction under poor soil moisture conditions. In this species, reproductive biomass and seed yield were proportional to current biomass gain and, in contrast to B. desertorum, independent of initial bulb mass, provided that the bulb was large enough to initiate flowering.

Life histories of desert geophytes -the demographic consequences of reproductive biomass partitioning patterns.

During five consecutive growing seasons (winters) ca. 110 plants of the desert geophyte Bellevalia desertorum were marked individually in a 25 m2 plot on a south-facing slope in the central Negev Desert of Israel. The number of rosette leaves of each plant was recorded, as well as whether it flowered and produced seeds. Multiple regression analysis of the data on the B. desertorum individuals showed that the reproductive state of a plant was determined by 1) its previous size and 2) the current conditions (rainfall), but not by previous conditions, nor by previous reproductive activity. Plant surveys supported these findings. These demographic results were consistent with the current understanding of the reproductive resource allocation pattern of B. desertorum. Flowering was most affected by rainfall until January of the same season, the number of leaves by rainfall until March and seed set was by the total annual amount. Surveys in populations ofB. eigii, a species of more mesic habitats in the Negev Desert suggested that in this species there is a negative effect of previous reproduction in combination with the previous and current conditions, which is also expected from its biomass partitioning pattern.The significance of demographic studies of individual plants in natural populations for the interpretation of experimentally determined resource partitioning patterns was discussed.