Vegetation spatial patterns and their effects on ecosystem functioning

-------In the Patagonian steppe, vegetation is composed of shrubs and tussock grasses. They are arranged in two patch types: shrubs encircled by a dense ring of tussocks grasses and tussock grasses interspersed with bare soil (Soriano et al. 1994). A model has been proposed for the origin and maintenance of this patchy structure in which spatial heterogeneity is viewed as reflecting phases of a cyclical succession (Fig. 3). When a shrub seedling establishes, it grows, and after achieving a certain size generates favorable conditions for grass establishment. These favorable conditions result in the construction of a dense ring of grasses surrounding the shrub. The building of the ring represents the construction phase of the cycle. When the shrub dies and aerial protection disappears, the density of grasses becomes higher than the current carrying capacity of the ring that results in the death of some tussocks. The ring disappears as an entity, and grasses appear interspersed with bare soil patches forming the other patch type of the Patagonian steppe. This would be the destruction phase of the cycle.

A heterogeneous vegetation structure results in the concentration of resources underneath individual plants, patches, or stripes. Two decades ago, Charley and West, (1975) introduced the term islands of fertility to describe the accumulation of nutrients underneath the canopy of shrubs in the semi-desert of Utah. Similarly, in the Chilean matorral, the concentration of soil nitrogen, phosphorus, and soil organic matter were higher underneath than outside the canopies of shrubs (Gutierrez et al. 1993). Smaller individuals such as those of the grass species Bouteloua gracilis were able to significantly modify the concentration of nitrogen and carbon underneath their canopies (Hook et al. 1991). At a coarser scale, the upper soil layer of the mulga groves had higher available nitrogen than the inter-groves (Ludwig and Tongway 1995). Phosphorus being an element less mobile in the soil showed similar trends but only in the uppermost layer.

What is the effect of this heterogeneous distribution of resources on the functioning of the entire ecosystem? Does it enhance or reduce production? Does it enhance or reduce ecosystem functioning in general? The answers to these questions have profound theoretical and applied implications. They can influence our understanding of the functioning of arid rangelands and our selection of management techniques that may try to either avoid or take advantage of heterogeneity. Schlesinger, et al. (1990) suggested that heterogeneity in the distribution of resources was a result of desertification while others indicate that is necessary for the survival of many plants and animals in arid and semiarid rangelands (Stafford Smith and Pickup 1990).

-------We suggest that the model that relates production and precipitation is linear and has two thresholds (Fig. 4A). Below a minimum precipitation threshold production is zero, and beyond a maximum precipitation threshold further increases in precipitation do not result in increases in production. There is empirical support of the existence of the minimum and maximum thresholds. Analysis of the production patterns of 9498 grassland sites in the Central Grassland Region of North America indicated that primary production and precipitation are positively correlated and the linear model has a negative y-intercept (Sala et al. 1988): 

            ANPP=-34 + 0.6*APPT

where ANPP is aboveground net primary production and APPT is annual precipitation. This equation can be rearranged as:

            ANPP= 0.6*(APPT-56)

where 0.6 represents the average water use efficiency and 56 is the ineffective precipitation or zero-yield intercept (Noy-Meir 1973). The analysis of the production patterns of forests, grasslands, and deserts suggested the existence of an upper threshold in the response of production to precipitation (Webb et al. 1978, Webb et al. 1983).

            This model has important implications for the effect of spatial heterogeneity on total ecosystem production based on the idea that in a patchy environment there are areas which concentrate resources, sinks, and areas which provide resources, sources. In the case of water, those are run-on and run-off areas. The amount of water redistributed depends on the proportion of sources and sinks, and on the fraction of precipitation which runs off which in turn depends on the magnitude of the precipitation event, the soil water content at the time of the event, topography, plant cover, and stocking rate (Branson et al. 1981).  If average precipitation falls below the lower threshold and it is homogeneously distributed, available water is (Pa) and production should be zero (Fig. 4B). If there is redistribution, the source areas will give up some water and will end up with less available water (Po), and production will continue to be zero. However, the sink areas will receive precipitation plus run-on that will result in a higher water availability (Pi) which now is higher than the threshold. Consequently, there will be production in the sink areas. This analysis suggests that in arid regions heterogeneous systems may have higher production than homogeneous systems. On the contrary, in mesic grasslands in which average precipitation is close to the upper threshold, heterogeneous systems should have lower production than homogeneous systems. This analysis was performed for water redistribution and it can be similarly performed for some other resources that limit production at different points in time. Nitrogen and phosphorus can be redistributed as water. In the case of nutrients, biological mechanisms can be invoked in addition to abiotic mechanisms to account for the redistribution of nutrients. Roots forage for nutrients beyond canopy projection and most of the aboveground litter is deposited underneath the canopy. Finally, animals may contribute to the redistribution of nutrients and concentration in patches.

In this project, we address both the effect of land-use change (i.e., grazing) and the effect of climate change (i.e., precipitation) on the spatial pattern of arid ecosystems. Our main underlying objective is to identify how these two driving forces control the vegetation spatial patterns and how they affect the functioning of ecosystems. We approached this issue with a combination of observation and field manipulative experiments.

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References
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