Ecosystem responses to increased variability in rainfall, a prediction of general circulation models, were assessed in native grassland by reducing storm frequency and increasing rainfall quantity per storm during a 4-year experiment. More extreme rainfall patterns, without concurrent changes in total rainfall quantity, increased temporal variability in soil moisture and plant species diversity. However, carbon cycling processes such as soil CO2 flux, CO2 uptake by the dominant grasses, and aboveground net primary productivity (ANPP) were reduced, and ANPP was more responsive to soil moisture variability than to mean soil water content. Our results show that projected increases in rainfall variability can rapidly alter key carbon cycling processes and plant community composition, independent of changes in total precipitation.

Although the potential for increased temperature is the primary and best-studied aspect of anthropogenic climate change, altered rainfall patterns, increased storm intensity, and more severe droughts are also predicted in most climate-change scenarios. We altered experimentally the rainfall regime in a native tallgrass prairie in northeastern Kansas and assessed leaf-level physiological activity and plant growth responses for C 3 and C 4 plant species. Our primary objective was to contrast the importance of reductions in rainfall quantity (30% smaller rain events, no change in rainfall pattern) with an altered, more extreme distribution of rainfall (no reduction in total growing-season quantity, 50% increased inter-rainfall dry intervals) for these dominant species from the two main plant functional groups (C 4 grasses, C 3 forbs) present in many grasslands. Leaf water potential (w l), net photosynthetic carbon gain (), specific leaf mass, leaf C : N ratio, growth rate A CO2 for Andropogon gerardii (C 4 grass) and Solidago canadensis (C 3 forb), vegetative and flowering stem densities, and canopy light penetration for grass and forb assemblages were intensively monitored during the 1999 growing season in a long-term rainfall manipulation study at the Konza Prairie Biological Station. Soil water content at 0-30 cm depth was more variable in response to the altered rainfall distribution compared to the reduced-quantity treatment. In S. canadensis, , g s (stomatal conductance), A : Ci (leaf stomatal [CO 2 ]), A CO2 and A : E (estimated leaf transpiration rate) were positively correlated with soil water content, but no relationship was seen for A. gerardii, indicating that even though this dominant grass species has most of its roots in the upper 30 cm of the soil, A. gerardii was buffered physiologically from increased resource variability. There were few significant responses in growth parameters in either grasses or forbs, but canopy light penetration increased with both rainfall treatments. We concluded (1) that the temporal variability in rainfall inputs can have as much impact on soil moisture as simple reductions in rainfall quantities with no change in temporal distribution, (2) that responses of A. gerardii and S. canadensis to altered rainfall distributions were not consistent with common views of soil resource partitioning between shallow-rooted grasses and deep-rooted forbs, and (3) that altered rainfall patterns may have the potential to offset elevated CO 2 impacts on grassland vegetation.

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.

Seedling growth dynamics of Quercus macrocarpa Michx. and Quercus muhlenbergii Engelm. were compared over a 3-month period under optimal growth conditions. These two species are the dominant trees at the western limit of the eastern deciduous forest, and are typically confined to gallery forests along stream beds in tallgrass prairie. Since tallgrass prairie is characterized by a highly variable climate and is prone to periodic drought, we hypothesized that these oaks would have rapid root growth and produce deep taproots as seedlings, enabling them to avoid drought stress and persist in this region. These traits may also facilitate forest expansion into the more xeric tallgrass prairie if fires are suppressed. Taproots of Q. macrocarpa and Q. muhlenbergii grew to approximately 140 cm and 100 cm in length, respectively, after 104 days. In both species, 65% or more of seedling biomass was allocated below ground, and root/total biomass was significantly greater in Q. muhlenbergii at 0-20 and 21-40 days after germination. The seedling taproot elongation rates reported here are much greater than rates reported in other eastern deciduous forest trees. Long-term precipitation data and soil moisture patterns from tallgrass prairie, when combined with rapid taproot elongation rates, suggest that soil moisture may not limit oak establishment or growth in tallgrass prairie in most years, although water uptake by roots was not measured in this study. Other factors, such as fire, herbivory, and seed predation and dispersal may be equally important in constraining the distribution of these species to gallery forests.

Abstract. In ecosystems where environments are extreme, such as deserts, adult plant species may facilitate the establishment and growth of seedlings and juveniles. Because high temperatures and evaporative demand characterize tall‐grass prairies of the central United States (relative to forests), we predicted that the grassland‐forest ecotone, by minimizing temperature extremes and moderating water stress, may function to facilitate the expansion of Quercus species into undisturbed tall‐grass prairie. We assessed the carbon and water relations of juvenile Quercus macrocarpa and Q. muhlenbergii, the dominant tree species in gallery forests of northeast Kansas, in ecotone and prairie sites. To evaluate the potentially competitive effects of neighboring herbaceous biomass on these oaks, juveniles (< 0.5 m tall) of both species also were subjected to either: (1) removal of surrounding above‐ground herbaceous biomass, or (2) control (prairie community intact) treatments. Herbaceous biomass removal had no significant effect on gas exchange or water relations in these oak species in either the prairie or the ecotone environment. Although the ecotone did alleviate some environmental extremes, photosynthetic rates and stomatal conductance were ca. 20 % higher (p < 0.05) in both oaks in prairie sites vs. the ecotone. Moreover, although leaf temperatures on average were higher in oaks in the prairie, high leaf temperatures in the ecotone had a greater negative effect on photosynthesis. These data suggest that the grassland‐forest ecotone did not facilitate the growth of Quercus juveniles expanding into this grassland. Moreover, the carbon and water relations of juvenile oaks in the prairie appeared to be unaffected by the presence of the dominant C4 grasses.