Lara G. Reichmann

Legacies of precipitation fluctuations on primary production: theory and data synthesis

Variability of above-ground net primary production (ANPP) of arid to sub-humid ecosystems displays a closer association with precipitation when considered across space (based on multiyear averages for different locations) than through time (based on year-to-year change at single locations). Here, we propose a theory of controls of ANPP based on four hypotheses about legacies of wet and dry years that explains space versus time differences in ANPP–precipitation relationships. We tested the hypotheses using 16 long-term series of ANPP. We found that legacies revealed by the association of current- versus previous-year conditions through the temporal series occur across all ecosystem types from deserts to mesic grasslands. Therefore, previous-year precipitation and ANPP control a significant fraction of current-year production. We developed unified models for the controls of ANPP through space and time. The relative importance of current-versus previous-year precipitation changes along a gradient of mean annual precipitation with the importance of current-year PPT decreasing, whereas the importance of previous-year PPT remains constant as mean annual precipitation increases. Finally, our results suggest that ANPP will respond to climate-change-driven alterations in water availability and, more importantly, that the magnitude of the response will increase with time.

Precipitation legacies in desert grassland primary production occur through previous‐year tiller density

In arid ecosystems, current-year precipitation often explains only a small proportion of annual aboveground net primary production (ANPP). We hypothesized that lags in the response of ecosystems to changes in water availability explain this low explanatory power, and that lags result from legacies from transitions from dry to wet years or the reverse. We explored five hypotheses regarding the magnitude of legacies, two possible mechanisms, and the differential effect of previous dry or wet years on the legacy magnitude. We used a three-year manipulative experiment with five levels of rainfall in the first two years (-80% and -50% reduced annual precipitation (PPT), ambient, +50% and +80% increased PPT), and reversed treatments in year 3. Legacies of previous two years, which were dry or wet, accounted for a large fraction (20%) of interannual variability in production on year 3. Legacies in ANPP were similar in absolute value for both types of precipitation transitions, and their magnitude was a function of the difference between previous and current-year precipitation. Tiller density accounted for 40% of legacy variability, while nitrogen and carryover water availability showed no effect. Understanding responses to changes in interannual precipitation will assist in assessing ecosystem responses to climate change-induced increases in precipitation variability.

Water controls on nitrogen transformations and stocks in an arid ecosystem

Following water, nitrogen (N) is the most frequent limiting factor to aboveground net primary production in arid ecosystems. Increased water availability can stimulate both plant nitrogen uptake and microbial nitrogen mineralization, but may also stimulate losses from the ecosystem. Here, we assess the effect of water availability on nitrogen stocks and transformations in an arid ecosystem. We conducted a field experiment with five levels of precipitation input (−80%, −50%, ambient, +50%, +80%) and two levels of N fertilization (ambient or 10 g·m−2·yr−1 NH4NO3) in a desert grassland of the Chihuahuan Desert. We measured in situ net N mineralization, plant N uptake, foliar N, N leaching under grass-rooting zone, and soil N availability during two years. Our results showed that increased water availability did not affect net N mineralization, but there was higher plant N uptake than with drought. Soil inorganic N pools were 2–4 times lower with increased water availability compared to drought conditions. N leaching below grass-rooting zone was higher in dry than wet conditions because of higher available N. Increased water availability differentially affected N species significantly reducing the NO3:NH4 ratio. The accumulation of inorganic N during drought was the result of a decoupling between microbial and plant activity, and suggests that the cycling of N is more open in dry years than in wet years.

Soil animal responses to moisture availability are largely scale, not ecosystem dependent: insight from a cross-site study

Climate change will result in reduced soil water availability in much of the world either due to changes in precipitation or increased temperature and evapotranspiration. How communities of mites and nematodes may respond to changes in moisture availability is not well known, yet these organisms play important roles in decomposition and nutrient cycling processes. We determined how communities of these organisms respond to changes in moisture availability and whether common patterns occur along fine-scale gradients of soil moisture within four individual ecosystem types (mesic, xeric and arid grasslands and a polar desert) located in the western United States and Antarctica, as well as across a cross-ecosystem moisture gradient (CEMG) of all four ecosystems considered together. An elevation transect of three sampling plots was monitored within each ecosystem and soil samples were collected from these plots and from existing experimental precipitation manipulations within each ecosystem once in fall of 2009 and three times each in 2010 and 2011. Mites and nematodes were sorted to trophic groups and analyzed to determine community responses to changes in soil moisture availability. We found that while both mites and nematodes increased with available soil moisture across the CEMG, within individual ecosystems, increases in soil moisture resulted in decreases to nematode communities at all but the arid grassland ecosystem; mites showed no responses at any ecosystem. In addition, we found changes in proportional abundances of mite and nematode trophic groups as soil moisture increased within individual ecosystems, which may result in shifts within soil food webs with important consequences for ecosystem functioning. We suggest that communities of soil animals at local scales may respond predictably to changes in moisture availability regardless of ecosystem type but that additional factors, such as climate variability, vegetation composition, and soil properties may influence this relationship over larger scales.

Differential sensitivities of grassland structural components to changes in precipitation mediate productivity response in a desert ecosystem

Summary In arid and semi-arid ecosystems, there are legacies of previous-year precipitation on current-year above-ground net primary production. We hypothesized that legacies of past precipitation occur through changes in tiller density, stolon density, tiller growth, axillary bud density and percentage of viable axillary buds. We examined the sensitivity to current- and previous-year precipitation of these grassland structural components in Bouteloua eriopoda, the dominant grass in the northern Chihuahuan Desert. We conducted a rainfall manipulation experiment consisting in −80% reduced precipitation, ambient, +80% increased precipitation treatments that were subjected to one of five precipitation levels in the previous two years (−80% and −50% reduced precipitation, ambient, +50% and +80% increased precipitation). The first two years preconditioned the experimental plots for year three, in which we created wet-to-dry and dry-to-wet transitions. Measurements were taken in year 3. We found that stolon density was the most sensitive to changes in precipitation and that percent-active buds were insensitive. We also found that past precipitation had a significant legacy on grassland structural components regardless of the precipitation received in the current year, and that the legacy occurs mostly through changes in stolon density. Here, we showed that there is a differential sensitivity of structural components to current and past precipitation and supported previous findings that vegetation structure is one of the controls of productivity during precipitation transitions.

Climate modifies response of non-native and native species richness to nutrient enrichment

Ecosystem eutrophication often increases domination by non-natives and causes displacement of native taxa. However, variation in environmental conditions may affect the outcome of interactions between native and non-native taxa in environments where nutrient supply is elevated. We examined the interactive effects of eutrophication, climate variability and climate average conditions on the success of native and non-native plant species using experimental nutrient manipulations replicated at 32 grassland sites on four continents. We hypothesized that effects of nutrient addition would be greatest where climate was stable and benign, owing to reduced niche partitioning. We found that the abundance of non-native species increased with nutrient addition independent of climate; however, nutrient addition increased non-native species richness and decreased native species richness, with these effects dampened in warmer or wetter sites. Eutrophication also altered the time scale in which grassland invasion responded to climate, decreasing the importance of long-term climate and increasing that of annual climate. Thus, climatic conditions mediate the responses of native and non-native flora to nutrient enrichment. Our results suggest that the negative effect of nutrient addition on native abundance is decoupled from its effect on richness, and reduces the time scale of the links between climate and compositional change.

Responses of a desert nematode community to changes in water availability

The most recent climate models unequivocally predict a strong drying trend for the southwestern USA within the next century. Soil nematodes are a highly important component of desert ecosystem functioning, but rely on water films for movement. However, it is currently poorly understood how different trophic groups of nematodes respond to chronic presses as well as short-lived pulses of altered water availability, especially in arid systems where such changes are expected to have the greatest impact. The aim of this study was to assess the effects of both instantaneous and long-term variation in water availability on desert soil nematode trophic groups. We hypothesized that nematode abundance would respond positively to both short- and long-term increases in moisture. Based on the ecology of the different trophic groups we further made predictions about their relative rates of response. We increased or decreased precipitation from ambient levels in the Chihuahuan Desert for four consecutive years and sampl...

Accelerated development in Johnsongrass seedlings (Sorghum halepense) suppresses the growth of native grasses through size-asymmetric competition

Invasive plant species often dominate native species in competition, augmenting other potential advantages such as release from natural enemies. Resource pre-emption may be a particularly important mechanism for establishing dominance over competitors of the same functional type. We hypothesized that competitive success of an exotic grass against native grasses is mediated by establishing an early size advantage. We tested this prediction among four perennial C4 warm-season grasses: the exotic weed Johnsongrass (Sorghum halepense), big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparius) and switchgrass (Panicum virgatum). We predicted that a) the competitive effect of Johnsongrass on target species would be proportional to their initial biomass difference, b) competitive effect and response would be negatively correlated and c) soil fertility would have little effect on competitive relationships. In a greenhouse, plants of the four species were grown from seed either alone or with one Johnsongrass neighbor at two fertilizer levels and periodically harvested. The first two hypotheses were supported: The seedling biomass of single plants at first harvest (50 days after seeding) ranked the same way as the competitive effect of Johnsongrass on target species: Johnsongrass < big bluestem < little bluestem/switchgrass, while Johnsongrass responded more strongly to competition from Johnsongrass than from native species. At final harvest, native plants growing with Johnsongrass attained between 2–5% of their single-plant non-root biomass, while Johnsongrass growing with native species attained 89% of single-plant non-root biomass. Fertilization enhanced Johnsongrass’ competitive effects on native species, but added little to the already severe competitive suppression. Accelerated early growth of Johnsongrass seedlings relative to native seedlings appeared to enable subsequent resource pre-emption. Size-asymmetric competition and resource-pre-emption may be a critical mechanism by which exotic invasive species displace functionally similar native species and alter the functional dynamics of native communities.