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Whole-plant hydraulics provide important information about responses to water limitation and can be used to understand how plant communities may change in a drier climate when measured on multiple species. Here, we measured above- and belowground hydraulic traits in Cornus drummondii, an encroaching shrub within North American tallgrass prairies, and Andropogon gerardii, a dominant C4 grass, to assess the potential hydraulic responses to future drought as this region undergoes woody expansion. Shelters that reduced precipitation by 50% and 0% were built over shrubs and grasses growing in sites that are burned at 1-year and 4-year frequencies. We then measured aboveground (Kshoot), belowground (Kroot), and whole-plant maximum hydraulic conductance (Kplant) in C. drummondii and Kroot in A. gerardii. We also measured vulnerability to embolism (P50) in C. drummondii stems. Overall, we show that: (1) A. gerardii had substantially greater Kroot than C. drummondii; (2) belowground hydraulic functioning was linked with aboveground processes; (3) above- and belowground C. drummondii hydraulics were not negatively impacted by the rainfall reductions imposed here. These results suggest that a multi-year drought will not ameliorate rates of woody expansion and highlight key differences in aboveground and belowground hydraulics for dominant species within the same ecosystem. [ABSTRACT FROM AUTHOR]

Ecosystems are faced with an onslaught of co-occurring global change drivers. While frequently studied independently, the effects of multiple global change drivers have the potential to be additive, antagonistic, or synergistic. Global warming, for example, may intensify the effects of more variable precipitation regimes with warmer temperatures increasing evapotranspiration and thereby amplifying the effect of already dry soils. Here, we present the long-term effects (11 years) of altered precipitation patterns (increased intra-annual variability in the growing season) and warming (1 °C year-round) on plant community composition and aboveground net primary productivity (ANPP), a key measure of ecosystem functioning in mesic tallgrass prairie. Based on past results, we expected that increased precipitation variability and warming would have additive effects on both community composition and ANPP. Increased precipitation variability altered plant community composition and increased richness, with no effect on ANPP. In contrast, warming decreased ANPP via reduction in grass stems and biomass but had no effect on the plant community. Contrary to expectations, across all measured variables, precipitation and warming treatments had no interactive effects. While treatment interactions did not occur, each treatment did individually impact a different component of the ecosystem (i.e., community vs. function). Thus, different aspects of the ecosystem may be sensitive to different global change drivers in mesic grassland ecosystems. [ABSTRACT FROM AUTHOR]

Increasing nitrogen deposition threatens many grassland species with local extinction. In addition to the direct effects of nitrogen deposition, nitrogen can indirectly affect plant populations via phenotypic shifts in plant traits that influence plant susceptibility to herbivory. Here, I test how herbivory varies across an experimental nitrogen gradient and whether differences in susceptibility to herbivory might explain patterns of local species loss. Specifically, I examine how increasing nitrogen availability in a restored prairie influences leaf traits and subsequent herbivory (by leaf-chewers like insects/small mammals versus deer) and the severity of herbivore damage on confamiliar pairs of extirpated versus extant species from Michigan prairies. Nitrogen increased herbivory by both leaf-chewers and deer as well as herbivore damage (proportion of leaves damaged). Leaf hairiness and specific leaf area affected patterns of herbivory following nitrogen addition, although patterns varied between extirpated vs. extant taxa and herbivory type. Nitrogen increased leaf hairiness. At high levels of nitrogen addition, hairy extant plants experienced less herbivory and damage than smooth-leaved plants. In contrast, hairy extirpated plants were more likely to experience leaf-chewer herbivory. Extirpated plants with thin leaves (high specific leaf area) were less likely to experience leaf-chewer herbivory; the opposite was true for extant species. Generally, extant species experienced more herbivory than locally extirpated species, particularly at high levels of nitrogen addition, suggesting that increasing herbivory under nutrient addition likely does not influence extirpation in this system. This study suggests that trait-mediated responses to nitrogen addition and herbivory differ between extant and extirpated species. [ABSTRACT FROM AUTHOR]

Woody encroachment—the spread of woody vegetation in open ecosystems—is a common threat to grasslands worldwide. Reversing encroachment can be exceedingly difficult once shrubs become established, particularly clonal species that resprout following disturbance. Single stressors are unlikely to reverse woody encroachment, but using multiple stressors in tandem could be successful in slowing or reversing encroachment. We explored whether increasing fire frequency in conjunction with multi-year drought could reduce growth and survival of encroaching shrubs in a tallgrass prairie in northeastern Kansas, USA. Passive rainout shelters (~ 50% rainfall reduction) were constructed over mature clonal shrubs (Cornus drummondii) and co-existing C4 grasses in two fire treatments (1-year and 4-year burn frequency). Leaf- and whole-plant level physiological responses to drought and fire frequency were monitored in shrubs and grasses from 2019 to 2022. Shrub biomass and stem density following fire were unaffected by five years of consecutive drought treatment, regardless of fire frequency. The drought treatment had more negative effects on grass leaf water potential and photosynthetic rates compared to shrubs. Shrub photosynthetic rates were remarkably stable across each growing season. Overall, we found that five consecutive years of moderate drought in combination with fire was not sufficient to reduce biomass production or stem density in an encroaching clonal shrub (C. drummondii). These results suggest that moderate but chronic press-drought events do not sufficiently stress encroaching clonal shrubs to negatively impact their resilience following fire events, even when fire frequency is high. [ABSTRACT FROM AUTHOR]

Cover crops are increasingly being used in ecological restoration projects, and are hypothesized to facilitate establishment of sown species by reducing weed abundances without competing with the target mix. We tested these predictions and examined the role of cover crop species on later species composition and diversity using cover crop seed treatments. Treatments included a fall seeding of one annual (Raphanus sativus or Avena sativa), one biennial (Oenothera biennis), one perennial species (Elymus canadensis), two grass-forb species combinations, or nothing as a control. All plots received the same diverse tallgrass prairie seed mix in March of the following year. Plant communities were sampled through five growing seasons. We found that cover crop treatments influenced community assembly, and that cover crop species varied in their effectiveness at reducing weed abundances, with the perennial grass E. canadensis being especially effective at reducing weeds. After 5 years, treatments were neutral in their effects on species diversity. However, composition of establishing plants (not including the cover crop) differed significantly among cover crop treatments on all sampling dates, indicating that treatments had long-term effects. Plots containing the C3 grass E. canadensis had greater abundance of C3 grasses and forbs, and reduced C4 grass abundance compared to other treatments. Our results indicate that cover crop species differ in their effects on assembling tallgrass prairie communities and that niche modification (i.e., species altering the abiotic environment in a way that favors species that benefit from the alterations) occurs when the cover crop is a perennial grass. [ABSTRACT FROM AUTHOR]

Root hemiparasites acquire resources from neighboring plants' vascular systems and can limit host growth, depress community productivity, and exert keystone effects. The strength of these effects is posited to be greater where hosts are nutrient-stressed but studies of annual hemiparasites show effects to be short-lived and variable. We conducted a 10-year experiment testing whether fertilizer addition alters the impact of the clonal, perennial hemiparasite Pedicularis canadensis on a prairie community and examine whether short-term trends reflect longer-term effects on community dynamics. Hemiparasite removal in 1-m2 plots increased productivity over the first three field seasons, but later the difference between removal and non-removal plots diminished as P. canadensis disappeared from 24 of the 48 non-removal plots. Effects of hemiparasite removal were context independent relative to fertilizer and shade treatments, but fertilizer initially increased, and then subsequently suppressed P. canadensis biomass. In non-removal plots, hemiparasite biomass was negatively associated with total community dry mass, which was greater in fertilized plots. Initially, fertilizer promoted graminoids, but after seven more field seasons, non-legume forbs responded most strongly. Measures of biodiversity tended to increase with hemiparasite cover. Demographic data collected at two different times for P. canadensis show high survivorship of established plants, high seed input, with seedling survival greater in taller vegetation. Unlike annual hemiparasite populations, well-established P. canadensis buffer populations against large demographic swings. At the scale of a few square meters, this keystone species produces significant heterogeneity in a prairie, but its presence at that scale is transient over approximately one decade. [ABSTRACT FROM AUTHOR]

Phylogenetic ecology uses evolutionary history to improve understanding of plant interactions. Phylogenetic distance can mediate plant interactions such as competition (e.g., via limiting similarity) and facilitation (e.g., via niche complementarity), influencing community assembly patterns. Previous research has found evidence both for and against a relationship between phylogenetic distance and the strength of plant interactions, and has found that other factors, such as trait differences, may be more influential. In addition to phylogenetic distance and species' traits, environmental conditions can also influence competition, with facilitative interactions—particularly among distantly related species—potentially becoming more pronounced under stressful, resource-limited conditions. We tested the prediction that greater phylogenetic distance is associated with decreased competition in a greenhouse experiment using plant species of the North American tallgrass prairie. We calculated the Relative Interaction Index for 81 species pairs using plant height, leaf length, and biomass as indicators of performance. We found that phylogenetic distance alone did not significantly affect competition. However, the interaction between phylogenetic distance and stressful conditions (sandier soils with low nutrient availability and water retention vs. resource-rich potting soil) altered plant traits and competition. Under stressful conditions, more distantly related species competed more strongly, leading to smaller plants. Conversely, under benign conditions more distantly related species pairs competed less and were larger. These results were contrary to our expectations that distant relatives would compete less under stressful conditions. Our experiment provides evidence that, while relatedness alone may not drive competition, phylogenetic distance can nonetheless be influential through interactions with environmental conditions. [ABSTRACT FROM AUTHOR]

Soil respiration, the major pathway for ecosystem carbon (C) loss, has the potential to enter a positive feedback loop with the atmospheric CO2 due to climate warming. For reliable projections of climate-carbon feedbacks, accurate quantification of soil respiration and identification of mechanisms that control its variability are essential. Process-based models simulate soil respiration as functions of belowground C input, organic matter quality, and sensitivity to environmental conditions. However, evaluation and calibration of process-based models against the long-term in situ measurements are rare. Here, we evaluate the performance of the Terrestrial ECOsystem (TECO) model in simulating total and heterotrophic soil respiration measured during a 16-year warming experiment in a mixed-grass prairie; calibrate model parameters against these and other measurements collected during the experiment; and explore whether the mechanisms of C dynamics have changed over the years. Calibrating model parameters against observations of individual years substantially improved model performance in comparison to pre-calibration simulations, explaining 79–86% of variability in observed soil respiration. Interannual variation of the calibrated model parameters indicated increasing recalcitrance of soil C and changing environmental sensitivity of microbes. Overall, we found that (1) soil organic C became more recalcitrant in intact soil compared to root-free soil; (2) warming offset the effects of increasing C recalcitrance in intact soil and changed microbial sensitivity to moisture conditions. These findings indicate that soil respiration may decrease in the future due to C quality, but this decrease may be offset by warming-induced changes in C cycling mechanisms and their responses to moisture conditions. [ABSTRACT FROM AUTHOR]

The expansion of woody species into grasslands has altered community structure and ecosystem function of grasslands worldwide. In tallgrass prairie of the Central Great Plains, USA, decreased fire frequency and intensity have increased the cover and abundance of woody species. In particular, clonal shrub cover has increased at accelerated rates due to vegetative reproduction and resprouting after disturbance. We measured the intra-clonal stem demography and relative growth rates (estimated change in woody biomass) of the shrub Cornus drummondii in response to fire frequency (4 vs 20 year burn intervals) and simulated browsing during the 2018 and 2019 growing seasons at Konza Prairie Biological Station (Manhattan, Kansas). Overall, infrequent fire (4 year burn interval) increased intra-clonal stem relative growth rates and shrub relative growth rates. Intra-clonal stem relative growth rates were reduced in unbrowsed clones in 2018 due to drought and simulated browsing reduced intra-clonal stem relative growth rates in 2019. Additionally, simulated browsing nearly eliminated flower production within clones but did not affect intra-clonal stem mortality or recruitment within a growing season. Fire in conjunction with simulated browsing reduced estimated relative growth rates for entire shrub clones. Browsed shrubs that experienced prescribed fire in 2017 had reduced intra-clonal stem densities compared to unbrowsed shrubs and stem densities of browsed shrubs did not recover in 2018 or 2019. These results illustrate that infrequent fire alone promotes the expansion of clonal shrubs in tallgrass prairie and multiple interacting disturbances (e.g., fire and browsing) are required to control the spread of clonal shrubs into grasslands. [ABSTRACT FROM AUTHOR]

In grasslands worldwide, modified fire cycles are accelerating herbaceous species extinctions. Fire may avert population declines by increasing survival, reproduction, or both. Survival and growth after fires may be promoted by removal of competitors or biomass and increasing resource availability. Fire-stimulated reproduction may also contribute to population growth through bolstered recruitment. We quantified these influences of fire on population dynamics in Echinacea angustifolia, a perennial forb in North American tallgrass prairie. We first used four datasets, 7–21 years long, to estimate fire's influences on survival, flowering, and recruitment. We then used matrix projection models to estimate growth rates across several burn frequencies in five populations, each with one to four burns over 15 years. Finally, we estimated the contribution of fire-induced changes in each vital rate to changes in population growth. Population growth rates generally increased with burning. The demographic process underpinning these increases depended on juvenile survival. In populations with high juvenile survival, fire-induced increases in seedling recruitment and juvenile survival enhanced population growth. However, in populations with low juvenile survival, small changes in adult survival drove growth rate changes. Regardless of burn frequencies, our models suggest populations are declining and that recruitment and juvenile survival critically influence population response to fire. However, crucially, increased seedling recruitment only increases population growth rates when enough new recruits reach reproductive maturity. The importance of recruitment and juvenile survival is especially relevant for small populations in fragmented habitats subject to mate-limiting Allee effects and inbreeding depression, which reduce recruitment and survival, respectively. [ABSTRACT FROM AUTHOR]

Root distribution patterns in soil are critical to understanding the interactions between climate and vegetation. However, it is not clear how climate change and land use practices affect belowground net primary productivity (BNPP) at various soil depths. In order to explore the effects of warming and clipping on root-distribution patterns along soil profile (0-15, 15-30, and 30-45 cm), we conducted a field experiment from 2005 to 2010 in a tallgrass prairie. We used infrared heaters to elevate soil temperature by approximately 2 °C and annual clipping to mimic hay harvest. Results showed that roots were not evenly distributed through the soil profile. On average across treatments and years, 53 and 83 % of the BNPP to 45 cm was distributed in the top 15- and 30-cm soil layers, respectively. Warming- and clipping-induced increases in BNPP were distributed to different soil depths at the proportions similar to those of BNPP. The proportional distribution of BNPP at various soil depths to total BNPP (0-45 cm) was little affected by warming, clipping, and their interactions, resulting in non-significant changes in the distribution of BNPP through the soil profile. These findings suggest that the proportionally vertical distribution of BNPP may remain stable even when the amount of BNPP changes simultaneously in response to climate change and land use practices. [ABSTRACT FROM AUTHOR]

Few field studies of natural populations have examined the factors influencing local extinctions and colonization of empty habitat patches for a prey species and its predator. In this study, I carried out a census of planthopper (Prokelisia crocea; Hemiptera: Delphacidae) and egg parasitoid (Anagrus columbi; Hymenoptera: Mymaridae) incidence and densities in 147 host-plant patches (Spartina pectinata; Poaceae) over seven planthopper generations in a tall-grass prairie landscape. For both species, the likelihood of going extinct in a patch was related to a number of patch-specific variables: density, temporal variability in density, proportion of hosts parasitized (planthopper only), host-plant density, patch size, patch isolation, and composition of the surrounding matrix. Colonization likelihood was related only to the physical attributes of the patch. There was high patch turnover in this prairie landscape. On average, planthoppers went extinct in 23% of the patches and A. columbi went extinct in 51% of the patches in each generation. For the planthopper, extinction likelihood increased with a decrease in patch size and the proportion of the matrix composed of mudflat. Parasitism of eggs had no effect on the extinction likelihood of local P. crocea populations, suggesting that A. columbi may not play a major role in the patch dynamics of its host. The likelihood of extinction for A. columbi was dependent on factors that spanned three trophic levels. An increase in plant density, decrease in host density and decrease in parasitoid density all increased the likelihood of A. columbi extinction within a patch. The dependency on multiple trophic levels may explain the higher extinction risk for the parasitoid than its host. A. columbi extinction was also affected by the matrix habitat surrounding the patch—the effect was the opposite of that for P. crocea. Finally, vacant patches were colonized at rates of 53% and 34% per generation for the planthopper and parasitoid, respectively. For both species, colonization probabilities decreased with an increase in patch isolation. High host densities in a patch also favored high rates of colonization by A. columbi. I discuss how anthropogenic changes to the prairie landscape can affect the metapopulation dynamics and persistence time of this host-parasitoid interaction. [ABSTRACT FROM AUTHOR]

Long-term soil carbon (C) storage is essential for reducing CO2 in the atmosphere. Converting unproductive and environmentally sensitive agricultural lands to grasslands for bioenergy production may enhance C storage. However, a better understanding of the interacting effects of grass functional composition (i.e., relative abundance of C4 and C3 grass cover) and soil productivity on C storage will help guide sustainable grassland management. Our objective was to examine the relationship between grass functional composition and potential C storage and how it varies with potential soil productivity. We estimated C inputs from above- and belowground net primary productivity (ANPP and BNPP), and heterotrophic respiration (RH) to calculate net ecosystem production (NEP), a measure of potential soil C storage, in grassland plots of relatively high- and low-productivity soils spanning a gradient in the ratio of C4 to C3 grass cover (C4:C3). NEP increased with increasing C4:C3, but only in potentially productive soils. The positive relationship likely stemmed from increased ANPP, rather than BNPP, which was possibly related to efficient resource-use and physiological/anatomical advantages of C4 plants. RH was negatively correlated with C4:C3, possibly because of changes in microclimate or plant–microbe interactions. It is possible that in potentially productive soils, C storage can be enhanced by favoring C4 over C3 grasses through increased ANPP and BNPP and reduced RH. Results also suggest that potential C storage gains from C4 productivity would not be undermined by a corresponding increase in RH. [ABSTRACT FROM AUTHOR]