@article {KNZ001983, title = {Shifts in plant functional composition following long-term drought in grasslands}, journal = {Journal of Ecology}, volume = {107}, year = {2019}, pages = {2133 - 2148}, abstract = {

\ 1. Plant traits can provide unique insights into plant performance at the community scale. Functional composition, defined by both functional diversity and community-weighted trait means (CWMs), can affect the stability of above-ground net primary production (ANPP) in response to climate extremes. Further complexity arises, however, when functional composition itself responds to environmental change. The duration of climate extremes, such as drought, is expected to increase with rising global temperatures; thus, understanding the impacts of long-term drought on functional composition and the corresponding effect that has on ecosystem function could improve predictions of ecosystem sensitivity to climate change.
\ 2. We experimentally reduced growing season precipitation by 66\% across six temperate grasslands for 4 years and measured changes in three indices of functional diversity (functional dispersion, richness and evenness), community-weighted trait means and phylogenetic diversity (PD). Specific leaf area (SLA), leaf nitrogen content (LNC) and (at most sites) leaf turgor loss point (πTLP) were measured for species cumulatively representing ~90\% plant cover at each site.
\ 3. Long-term drought led to increased community functional dispersion in three sites, with negligible effects on the remaining sites. Species re-ordering following the mortality/senescence of dominant species was the main driver of increased functional dispersion. The response of functional diversity was not consistently matched by changes in phylogenetic diversity. Community-level drought strategies (assessed as CWMs) largely shifted from drought tolerance to drought avoidance and/or escape strategies, as evidenced by higher community-weighted πTLP, SLA and LNC. Lastly, ecosystem drought sensitivity (i.e. relative reduction in ANPP in drought plots) was positively correlated with community-weighted SLA and negatively correlated with functional diversity.
\ 4. Synthesis. Increased functional diversity following long-term drought may stabilize ecosystem functioning in response to future drought. However, shifts in community-scale drought strategies may increase ecosystem drought sensitivity, depending on the nature and timing of drought. Thus, our results highlight the importance of considering both functional diversity and abundance-weighted traits means of plant communities as their collective effect may either stabilize or enhance ecosystem sensitivity to drought.

}, keywords = {LTER-KNZ, ANPP, Climate change, community weighted traits, Drought, Functional diversity, plant functional traits}, doi = {10.1111/1365-2745.13252}, url = {https://besjournals.onlinelibrary.wiley.com/doi/10.1111/1365-2745.13252}, author = {Griffin-Nolan, Robert J. and D.M. Blumenthal and Scott. L. Collins and Farkas, Timothy E. and Hoffman, Ava M. and Mueller, Kevin E. and Ocheltree, Troy W. and M.D. Smith and Whitney, Kenneth D. and Alan K. Knapp}, editor = {Jones, Holly} } @article {KNZ001631, title = {Differential sensitivity to regional-scale drought in six central US grasslands}, journal = {Oecologia}, volume = {177}, year = {2015}, pages = {949 -957}, abstract = {

Terrestrial ecosystems often vary dramatically in their responses to drought, but the reasons for this are unclear. With climate change forecasts for more frequent and extensive drought in the future, a more complete understanding of the mechanisms that determine differential ecosystem sensitivity to drought is needed. In 2012, the Central US experienced the fourth largest drought in a century, with a regional-scale 40 \% reduction in growing season precipitation affecting ecosystems ranging from desert grassland to mesic tallgrass prairie. This provided an opportunity to assess ecosystem sensitivity to a drought of common magnitude in six native grasslands. We tested the prediction that drought sensitivity is inversely related to mean annual precipitation (MAP) by quantifying reductions in aboveground net primary production (ANPP). Long-term ANPP data available for each site (mean length = 16 years) were used as a baseline for calculating reductions in ANPP, and drought sensitivity was estimated as the reduction in ANPP per millimeter reduction in precipitation. Arid grasslands were the most sensitive to drought, but drought responses and sensitivity varied by more than twofold among the six grasslands, despite all sites experiencing 40 \% reductions in growing season precipitation. Although drought sensitivity generally decreased with increasing MAP as predicted, there was evidence that the identity and traits of the dominant species, as well as plant functional diversity, influenced sensitivity. A more comprehensive understanding of the mechanisms leading to differences in drought sensitivity will require multi-site manipulative experiments designed to assess both biotic and abiotic determinants of ecosystem sensitivity.

}, keywords = {LTER-KNZ, Climate change, Functional diversity, Long-term ecological research, Precipitation, Primary production}, doi = {10.1007/s00442-015-3233-6}, url = {https://link.springer.com/article/10.1007\%2Fs00442-015-3233-6}, author = {Alan K. Knapp and Carroll, C.J.W. and Denton, E.M. and Kimberly J. La Pierre and Scott. L. Collins and M.D. Smith} } @article {KNZ001532, title = {Rainfall variability has minimal effects on grassland recovery from repeated grazing}, journal = {Journal of Vegetation Science}, volume = {25}, year = {2014}, pages = {36 -44}, abstract = {

Question Mesic grasslands experience a complex disturbance regime including frequent fire, grazing by large ungulates and strong inter-annual climate variability. As a result of climate change, growing season precipitation regimes are predicted to become more variable, with larger event sizes and longer dry periods resulting in more temporally dynamic soil moisture regimes. Increased climate variability is likely to interact with other disturbances, such as grazing, in grassland ecosystems. We investigated the individual and combined effects of increased rainfall variability and grazing on plant community composition, structure and function in an annually burned, native tallgrass prairie. Our overarching question was: are grazing impacts modified under a more variable precipitation regime? Location Konza Prairie, Kansas, USA. Methods Plots were established within a long-term rainfall manipulation experiment in which larger, but less frequent, rain events were imposed during the growing season without altering the total rain amount. We then simulated intense grazing pressure during one growing season by repeatedly clipping all graminoids to 5 cm and monitored recovery over 3 yr. Results Neither grazing nor rainfall treatments affected species richness; however, grazing decreased total and grass above-ground net primary production (ANPP) and increased forb ANPP relative to ungrazed plots. Grass stem density recovered from intense grazing under ambient rainfall but did not fully recover, even after 2 yr in the altered rainfall treatment. Conclusions We found that increased rainfall variability had little effect on tallgrass prairie structure and function, while grazing had large effects. Grazing and increased rainfall variability interacted to suppress grass stem density and delayed recovery relative to controls. Although stem density was reduced, individual stem size increased, resulting in no net change in ANPP. This suggests that ANPP in grazed and ungrazed North American tallgrass prairie may be relatively resilient under more temporally variable precipitation regimes.

}, keywords = {LTER-KNZ, Annual net primary production, Climate change, Clipping, disturbance, diversity, Grass{\textendash}forb interaction, Konza, Precipitation, Stem density}, doi = {10.1111/jvs.12065}, url = {https://onlinelibrary.wiley.com/doi/abs/10.1111/jvs.12065}, author = {Koerner, S.E. and Scott. L. Collins and John M. Blair and Alan K. Knapp and M.D. Smith} } @booklet {KNZ001539, title = {Long-term trends in ecological systems: a basis for understanding responses to global change. USDA Agriculture Research Service Publication, Technical Bulletin 1931. Washington, D.C}, year = {2012}, abstract = {

Peters, D.P.C., C.M. Laney, A.E. Lugo, et al. 2013. Long-Term Trends in Ecological Systems: A Basis for Understanding Responses to Global Change. U.S. Department of Agriculture, Technical Bulletin Number 1931. The EcoTrends Editorial Committee sorted through vast amounts of historical and ongoing data from 50 ecological sites in the continental United States including Alaska, several islands, and Antarctica to present in a logical format the variables commonly collected. This report presents a subset of data and variables from these sites and illustrates through detailed examples the value of comparing longterm data from different ecosystem types. This work provides cross-site comparisons of ecological responses to global change drivers, as well as longterm trends in global change drivers and responses at site and continental scales. Site descriptions and detailed data also are provided in the appendix section.

}, keywords = {LTER-KNZ, atmospheric chemistry, Climate change, cross-site comparisons, disturbance, ecological response, ecology, ecosystem, EcoTrends, experimental forests, global change, human demography, human population growth, Long Term Ecological Research (LTER), long-term datasets, Precipitation, rangeland, rangeland research stations, surface water chemistry}, author = {Peters, D.P.C. and Laney, C.M. and Lugo, A.E. and Scott. L. Collins and Driscoll, C.T. and Groffman, P.M. and Grove, J.M. and Alan K. Knapp and Kratz, T.K. and Ohman, M.D. and Waide, R.B. and Yao, J.} } @article {KNZ001496, title = {Past, present, and future roles of long-term experiments in the LTER Network}, journal = {Bioscience}, volume = {62}, year = {2012}, pages = {377 -389}, abstract = {

The US National Science Foundation\–funded Long Term Ecological Research (LTER) Network supports a large (around 240) and diverse portfolio of long-term ecological experiments. Collectively, these long-term experiments have (a) provided unique insights into ecological patterns and processes, although such insight often became apparent only after many years of study; (b) influenced management and policy decisions; and (c) evolved into research platforms supporting studies and involving investigators who were not part of the original design. Furthermore, this suite of long-term experiments addresses, at the site level, all of the US National Research Council\&$\#$39;s Grand Challenges in Environmental Sciences. Despite these contributions, we argue that the scale and scope of global environmental change requires a more-coordinated multisite approach to long-term experiments. Ideally, such an approach would include a network of spatially extensive multifactor experiments, designed in collaboration with ecological modelers that would build on and extend the unique context provided by the LTER Network.

}, keywords = {LTER-KNZ, Climate change, global change, long-term research, LTER Network, multifactor experiments}, doi = {10.1525/bio.2012.62.4.9}, url = {https://academic.oup.com/bioscience/article/62/4/377/243762}, author = {Alan K. Knapp and M.D. Smith and Hobbie, S.E. and Scott. L. Collins and Fahey, T.J. and Hansen, G.J.A. and Landis, D.A. and Kimberly J. La Pierre and Melillo, J.M. and Seastedt, T.R. and Shaver, G.R. and Webster, J.R.} } @article {KNZ001187, title = {Consequences of more extreme precipitation regimes for terrestrial ecosystems}, journal = {BioScience}, volume = {58}, year = {2008}, pages = {811 -821}, abstract = {

mplification of the hydrological cycle as a consequence of global warming is forecast to lead to more extreme intra-annual precipitation regimes characterized by larger rainfall events and longer intervals between events. We present a conceptual framework, based on past investigations and ecological theory, for predicting the consequences of this underappreciated aspect of climate change. We consider a broad range of terrestrial ecosystems that vary in their overall water balance. More extreme rainfall regimes are expected to increase the duration and severity of soil water stress in mesic ecosystems as intervals between rainfall events increase. In contrast, xeric ecosystems may exhibit the opposite response to extreme events. Larger but less frequent rainfall events may result in proportional reductions in evaporative losses in xeric systems, and thus may lead to greater soil water availability. Hydric (wetland) ecosystems are predicted to experience reduced periods of anoxia in response to prolonged intervals between rainfall events. Understanding these contingent effects of ecosystem water balance is necessary for predicting how more extreme precipitation regimes will modify ecosystem processes and alter interactions with related global change drivers.

}, keywords = {LTER-KNZ, Climate change, Drought, Ecosystems, Precipitation, soil water}, doi = {10.1641/B580908}, url = {https://academic.oup.com/bioscience/article/58/9/811/250853}, author = {Alan K. Knapp and Beier, C. and Briske, D.D. and Classen, A.T. and Luo, Y. and Reichstein, M. and M.D. Smith and Smith, S.D. and Bell, J.E. and Fay, P.A. and Heisler, J.L. and Leavitt, S.W and Sherry, R. and Smith, B. and Weng, E.} } @article {KNZ001203, title = {Increased precipitation event size increases aboveground net primary productivity in a semi-arid grassland}, journal = {Oecologia}, volume = {158}, year = {2008}, pages = {129 -140}, abstract = {

Water availability is the primary constraint to aboveground net primary productivity (ANPP) in many terrestrial biomes, and it is an ecosystem driver that will be strongly altered by future climate change. Global circulation models predict a shift in precipitation patterns to growing season rainfall events that are larger in size but fewer in number. This \“repackaging\” of rainfall into large events with long intervening dry intervals could be particularly important in semi-arid grasslands because it is in marked contrast to the frequent but small events that have historically defined this ecosystem. We investigated the effect of more extreme rainfall patterns on ANPP via the use of rainout shelters and paired this experimental manipulation with an investigation of long-term data for ANPP and precipitation. Experimental plots (n = 15) received the long-term (30-year) mean growing season precipitation quantity; however, this amount was distributed as 12, six, or four events applied manually according to seasonal patterns for May\–September. The long-term mean (1940\–2005) number of rain events in this shortgrass steppe was 14 events, with a minimum of nine events in years of average precipitation. Thus, our experimental treatments pushed this system beyond its recent historical range of variability. Plots receiving fewer, but larger rain events had the highest rates of ANPP (184 \± 38 g m\−2), compared to plots receiving more frequent rainfall (105 \± 24 g m\−2). ANPP in all experimental plots was greater than long-term mean ANPP for this system (97 g m\−2), which may be explained in part by the more even distribution of applied rain events. Soil moisture data indicated that larger events led to greater soil water content and likely permitted moisture penetration to deeper in the soil profile. These results indicate that semi-arid grasslands are capable of responding immediately and substantially to forecast shifts to more extreme precipitation patterns.

}, keywords = {LTER-KNZ, Climate change, Grasslands, Precipitation variability, Pulse-reserve paradigm, Rain event size}, doi = {10.1007/s00442-008-1116-9}, url = {https://link.springer.com/article/10.1007\%2Fs00442-008-1116-9}, author = {Heisler-White, J.L. and Alan K. Knapp and Kelly, E.F.} } @article {KNZ001070, title = {Growth responses of twodominant C4 grass species to altered water availability}, journal = {InternationalJournal of Plant Sciences}, volume = {167}, year = {2006}, pages = {1001 -1010}, abstract = {Identifying key ecophysiological traits that differ among dominant plant species and can be linked to species-specific responses to drought would improve our ability to forecast community and ecosystem responses to global climate change. The mesic grasslands of the central plains of North America are dominated by two C4 grass species, Andropogon gerardii and Sorghastrum nutans, which purportedly differ in their tolerance of water stress. Individuals of these two species were grown in the field under rain-out shelters and subjected to wet (watered every 2{\textendash}3 d) or dry (repeatedly subjected to wilting before watering) soil moisture regimes. A range of ecophysiological traits potentially important for tolerating water stress were concurrently measured. Although few traits differed between the species in the wet treatment, several traits were identified in the dry treatment that may enable A. gerardii to better tolerate drought. These were greater allocation to roots, reduced allocation to flowering, more rapid leaf turnover, and more rapid recovery of photosynthesis after wilting. The latter two traits may be particularly important for coping with increased variability in rainfall regimes in the future and are consistent with recently documented responses of A. gerardii to experimental increases in soil moisture variability.}, keywords = {LTER-KNZ, Andropogon gerardii, biomass allocation, Climate change, leaf turnover, photosynthesis, Sorghastrum nutans, water stress}, doi = {10.1086/505611}, author = {Swemmer, A.M. and Alan K. Knapp and M.D. Smith} } @article {KNZ00993, title = {Intra-annual rainfallvariability and grassland productivity: can the past predictthe future}, journal = {Plant Ecology}, volume = {184}, year = {2006}, pages = {65 -74}, abstract = {Precipitation quantity has been shown to influence grassland aboveground net primary productivity (ANPP) positively whereas experimental increases in of temporal variability in water availability commonly exhibit a negative relationship with ANPP. We evaluated long term ANPP datasets from the Konza Prairie Long Term Ecological Research (LTER) program (1984{\textendash}1999) to determine if similar relationships could be identified based on patterns of natural variability (magnitude and timing) in precipitation. ANPP data were analyzed from annually burned sites in native mesic grassland and productivity was partitioned into graminoid (principally C4 grasses) and forb (C3 herbaceous) components. Although growing season precipitation amount was the best single predictor of total and grass ANPP (r 2=0.62), several measures of precipitation variability were also significantly and positively correlated with productivity, independent of precipitation amount. These included soil moisture variability, expressed as CV, for June (r 2=0.45) and the mean change in soil moisture between weekly sampling periods in June and August (\%wv) (r 2=0.27 and 0.32). In contrast, no significant relationships were found between forb productivity and any of the precipitation variables (p>0.05). A multiple regression model combining precipitation amount and both measures of soil moisture variability substantially increased the fit with productivity (r 2=0.82). These results were not entirely consistent with those of short-term manipulative experiments in the same grassland, however, because soil moisture variability was often positively, not negatively related to ANPP. Differences in results between long and short term experiments may be due to low variability in the historic precipitation record compared to that imposed experimentally as experimental levels of variability exceeded the natural variability of this dataset by a factor of two. Thus, forecasts of ecosystem responses to climate change (i.e. increased climatic variability), based on data constrained by natural and recent historical rainfall patterns may be inadequate for assessing climate change scenarios if precipitation variability in the future is expected to exceed current levels.}, keywords = {LTER-KNZ, ANPP, Climate change, grassland, Precipitation variability, soil moisture, tallgrass prairie}, doi = {10.1007/s11258-005-9052-9}, author = {Jesse B. Nippert and Alan K. Knapp and J. M. Briggs} } @article {KNZ00863, title = {Productivity responses to altered rainfall patterns in a C4-dominated grassland}, journal = {Oecologia}, volume = {137}, year = {2003}, pages = {245 -251}, abstract = {Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50\%, reducing rainfall quantities by 30\%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0{\textendash}30 cm depth), a ~10\% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C4 grasses while productivity of the dominant C4 grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30\% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.}, keywords = {LTER-KNZ, Climate change, Konza Prairie, Net primary productivity, Precipitation, soil moisture}, doi = {10.1007/s00442-003-1331-3}, author = {P. Fay and Carlisle, J.D. and Alan K. Knapp and John M. Blair and Scott. L. Collins} } @article {KNZ00858, title = {Altering rainfall timing and quantity in a mesic grassland ecosystem: Design and performance of rainfall manipulation shelters}, journal = {Ecosystems}, volume = {3}, year = {2000}, pages = {308 -319}, abstract = {Global climate change is predicted to alter growing season rainfall patterns, potentially reducing total amounts of growing season precipitation and redistributing rainfall into fewer but larger individual events. Such changes may affect numerous soil, plant, and ecosystem properties in grasslands and ultimately impact their productivity and biological diversity. Rainout shelters are useful tools for experimental manipulations of rainfall patterns, and permanent fixed-location shelters were established in 1997 to conduct the Rainfall Manipulation Plot study in a mesic tallgrass prairie ecosystem in northeastern Kansas. Twelve 9 x 14{\textendash}m fixed-location rainfall manipulation shelters were constructed to impose factorial combinations of 30\% reduced rainfall quantity and 50\% greater interrainfall dry periods on 6 x 6{\textendash}m plots, to examine how altered rainfall regimes may affect plant species composition, nutrient cycling, and above- and belowground plant growth dynamics. The shelters provided complete control of growing season rainfall patterns, whereas effects on photosynthetic photon flux density, nighttime net radiation, and soil temperature generally were comparable to other similar shelter designs. Soil and plant responses to the first growing season of rainfall manipulations (1998) suggested that the interval between rainfall events may be a primary driver in grassland ecosystem responses to altered rainfall patterns. Aboveground net primary productivity, soil CO2 flux, and flowering duration were reduced by the increased interrainfall intervals and were mostly unaffected by reduced rainfall quantity. The timing of rainfall events and resulting temporal patterns of soil moisture relative to critical times for microbial activity, biomass accumulation, plant life histories, and other ecological properties may regulate longer-term responses to altered rainfall patterns.}, keywords = {LTER-KNZ, Climate change, floristic diversity, Grasslands, Konza Prairie, life histories, long-term research, Net primary production, precipitation patterns, rainout shelters, soil moisture}, doi = {10.1007/s100210000028}, author = {Fay, P.A. and Carlisle, J.D. and Alan K. Knapp and John M. Blair and Scott. L. Collins} } @article {KNZ00740, title = {Patterns and determinants of potential carbon gain in the C3 evergreen Yucca glauca (Liliaceae) in a C4 grassland}, journal = {American Journal of Botany}, volume = {87}, year = {2000}, pages = {230 -236}, abstract = {Yucca glauca is a C3 evergreen rosette species locally common in the C4-dominated grasslands of the central Great Plains. Most congeners of Y. glauca are found in deserts, and Y. glauca{\textquoteright}s morphological similarities to desert species (steeply angled leaves, evergreen habit) may be critical to its success in grasslands. We hypothesized that the evergreen habit of Y. glauca, coupled with its ability to remain physiologically active at cool temperatures, would allow this species to gain a substantial portion of its annual carbon budget when the C4 grasses are dormant. Leaf-level gas exchange was measured over an 18-mo period at Konza Prairie in northeast Kansas to assess the annual pattern of potential C gain. Two short-term experiments also were conducted in which nighttime temperatures were manipulated to assess the cold tolerance of this species. The annual pattern of C gain in Y. glauca was bimodal, with a spring productive period (maximum monthly photosynthetic rate = 21.1 {\textpm} 1.97 μmol{\textperiodcentered}m-2{\textperiodcentered}s-1) in March through June, a period of midseason photosynthetic depression, and a fall productive period in October (15.6 {\textpm} 1.25 μmol{\textperiodcentered}m-2{\textperiodcentered}s-1). The steeply angled leaves resulted in interception of photon flux density at levels above photosynthetic saturation throughout the year. Reduced photosynthetic rates in the summer may have been caused by low soil moisture, but temperature was strongly related (r2 = 0.37) to annual variations in photosynthesis, with nocturnal air temperatures below -5{\textdegree}C in the late fall and early spring, and high air temperatures (>32{\textdegree}C) in the summer, limiting gas exchange. Overall, 31\% of the potential annual carbon gain in Y. glauca occurred outside the {\textquotedblleft}frost-free{\textquotedblright} period (April{\textendash}October) at Konza Prairie and 43\% occurred when the dominant C4 grasses were dormant. Future climates that include warmer minimum temperatures in the spring and fall may enhance the success of Y. glauca relative to the C4 dominants in these grasslands.}, keywords = {LTER-KNZ, Climate change, cold tolerance, evergreen, Grasslands, Liliaceae, photosynthesis, tallgrass prairie, Water relations, Yucca}, url = {http://www.amjbot.org/content/87/2/230.short}, author = {Maragni, L.A. and Alan K. Knapp and McAllister, C.A.} }