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Native seed mixes are widely available to create pollinator habitat. One approach to understand the efficacy of pollinator seed mixes is to compare them to species found in high‐quality natural areas. Using plant species presence data from published lists, we compared weekly blooming richness between 196 seed mixes explicitly designed to support pollinators and 102 prairie remnants from eight states across the Midwest. Remnants had greater forb richness compared to mixes and had at least twice the blooming richness per week. Seed mixes included low proportions of species that bloomed in the spring (0–13% of species), whereas about 28% of remnant forbs bloomed before June. There was a sharp phenological blooming peak for mixes due to the vast majority of species blooming in late July through August, whereas remnants had greater phenological evenness across the growing season. We then investigated all observed early blooming plant species to identify the species best suited to alleviate the dissimilar spring blooming characteristics. We compiled 10 species characteristics that reflected species' feasibility for cultivation and restoration and blooming attributes, identifying a group of underutilized and commercially unavailable species. Pollinator seed mixes should facilitate blooming availability for the duration of the growing season to meet pollinator foraging requirements. [ABSTRACT FROM AUTHOR]

Species interaction effects on populations can vary in both magnitude (i.e. strong vs. weak) and sign (positive, negative, or no effect). Context‐dependent effects of species interactions occur when the sign or strength of the interaction's effect on population growth rate changes across abiotic gradients.We know that species can vary substantially in the degree of context dependence they exhibit, even across similar abiotic gradients. However, few studies have characterised context dependence of co‐occurring species, limiting our ability to understand the implications of context dependence for species interaction effects on community composition.Using over three decades of data collected for 13 tallgrass prairie forbs at the Konza Prairie Biological Station, we parameterise density structured population models that predict population dynamics as functions of abiotic conditions and bison herbivory. We use these models to estimate the degree of context dependence in responses to bison herbivory for 13 species across three abiotic gradients: weather, fire frequency and soil type.All species showed significant context dependence for fire frequency in the same direction, though with variable magnitude, such that herbivory increased cover with more frequent fires. Context dependence with weather and soil type varied dramatically across species in both direction and magnitude. For example, herbivory effects on 3/13 species were stronger in wet conditions, but herbivory effects on 5/13 species were stronger in dry conditions. Thus, context dependence exhibited by individual species, as opposed to effects of abiotic conditions on the relative abundances of species, could generate much of the weather‐dependent effects of herbivory on community composition.Synthesis: Our work suggests that species can vary dramatically in the presence, direction and magnitude of context dependence, even when occurring in the same community and when considering the same species interaction (i.e. response to a herbivore). In addition, we find that context dependence could drive substantial variation in the effect of species interactions on community characteristics (e.g. composition) across multiple abiotic gradients. [ABSTRACT FROM AUTHOR]

North American tallgrass prairie evolved with regular fire set by Indigenous people and grazing from large herbivores, including the plains bison (Bison bison). In contrast to uniform effects of prescribed fire, bison graze selectively, creating increased heterogeneity on the landscape. We examined plant community responses to reintroduced bison grazing in high-diversity native and restored tallgrass prairie in Illinois, USA. We evaluated plant community diversity and structure in paired grazed and ungrazed (fenced) plots. After 5 y of grazing at a relatively low target stocking rate of 1 animal unit per 5.2 ha (13 acres), there were no differences in plant community diversity, composition, nonnative:native species ratio, or visual obstruction readings between grazed and ungrazed plots. An interaction exists (or was found) between plant community type and grazing in which grazed savanna plots had greater grass relative abundance compared to ungrazed plots. Ongoing monitoring will evaluate long-term trends. [ABSTRACT FROM AUTHOR]

Grasslands are among the most imperilled ecosystems worldwide, and many have experienced degradation due to the loss of historical disturbance regimes and subsequent woody encroachment. Management practitioners often use physical and chemical management interventions in combination with fire to counter encroachment, altering aboveground structure and belowground function, respectively. This may disrupt the feedbacks that perpetuate encroachment and restore the herbaceous community.We use a large‐scale field experiment to assess the initial effects of different management interventions on woody vegetation persistence, abiotic habitat conditions, and herbaceous community composition. We evaluate these effects across seven sites spanning a natural soil moisture gradient to capture one aspect of environmental heterogeneity with which managers regularly contend.We found that chemical intervention, both with and without the addition of physical intervention, was most effective at reducing woody plant cover and abundance, and a second application reduced woody plant abundance by more than one application alone. We also found that any management intervention increased light availability and air temperature and decreased soil moisture, with the combination of physical and chemical interventions having the greatest effects. Finally, none of the management interventions affected herbaceous richness and functional group cover within the study period, indicating delayed or nonexistent effects on herbaceous community composition.Synthesis and application. Our findings suggest that management should focus on chemical intervention for the greatest effects on woody plant persistence and abiotic habitat conditions. Changes to herbaceous community composition may occur in the long term and seem likely since short‐term effects of management were successful in altering processes related to encroachment feedbacks. [ABSTRACT FROM AUTHOR]

Nitrogen (N) availability is a well‐known driver of ecosystem structure and function, but as air quality regulations continue to reduce atmospheric N deposition, there is a need to understand how managed and unmanaged ecosystems respond to widespread decreases in terrestrial N availability. Historical N eutrophication, from pollution or fertilisation, may continue to constrain contemporary responses to decreases in available N because of altered plant and microbial feedbacks. Thus, while certain management practices like prescribed fire remove N from grassland ecosystems, the role of fire supporting ecosystems recovering from chronic N input is unknown.To address this knowledge gap, we ceased a 30‐year N‐fertilisation treatment at a field experiment in a tallgrass prairie ecosystem crossed with burned and fire‐suppressed (unburned) treatments. We established subplots within each previously fertilised, recovering plot, fertilised at the same historical rate (10 g N m−2 year−1 as NH4NO3), to compare plant and soil properties in recovering plots with control (never‐fertilised) and still‐fertilised treatments within different fire regimes.We document different N‐fertilisation legacies among ecosystem properties in burned and unburned prairies recovering from N‐fertilisation. Soil N availability, nitrification and denitrification potentials in recovering plots remained higher than controls for 3–5 years—indicative of positive legacies—in both burned and unburned prairies, but burning did not reduce this legacy. In burned prairies, however, a positive legacy in above‐ground plant production persisted because a more productive grass species (switchgrass) replaced the previously dominant species (big bluestem) even though root C:N, but not soil C:N, increased to return back to control levels. Consequently, the main N loss pathways in burned and unburned prairies (pyrovolatilisation and microbially mediated processes, respectively) led to similar losses of soil total N (20–28 g N m−2) over 5 years.Synthesis: Our results indicate that N eutrophication induces positive legacies of ecosystem functions that can persist for at least half a decade. N‐induced legacies arise because of shifts in soil microbial N‐cycling and plant functional traits. As a result, different management practices may elicit similar trajectories of ecosystem recovery in terms of total and available soil N because of different plant and microbial feedbacks. [ABSTRACT FROM AUTHOR]

Enhancing resilience in formerly degraded ecosystems is an important goal of restoration ecology. However, evidence for the recovery of resilience and its underlying mechanisms require long‐term experiments and comparison with reference ecosystems. We used data from an experimental prairie restoration that featured long‐term soil heterogeneity manipulations and data from two long‐term experiments located in a comparable remnant (reference) prairie to (1) quantify the recovery of ecosystem functioning (i.e., productivity) relative to remnant prairie, (2) compare the resilience of restored and remnant prairies to a natural drought, and (3) test whether soil heterogeneity enhances resilience of restored prairie. We compared sensitivity and legacy effects between prairie types (remnant and restored) and among four prairie sites that included two remnant prairie sites and prairie restored under homogeneous and heterogeneous soil conditions. We measured sensitivity and resilience as the proportional change in aboveground net primary productivity (ANPP) during and following drought (sensitivity and legacy effects, respectively) relative to average ANPP based on 4 pre‐drought years (2014–2017). In nondrought years, total ANPP was similar between remnant and restored prairie, but remnant prairie had higher grass productivity and lower forb productivity compared with restored prairie. These ANPP patterns generally persisted during drought. The sensitivity of total ANPP to drought was similar between restored and remnant prairie, but grasses in the restored prairie were more sensitive to drought. Post‐drought legacy effects were more positive in the restored prairie, and we attributed this to the more positive and less variable legacy response of forb ANPP in the restored prairie, especially in the heterogeneous soil treatment. Our results suggest that productivity recovers in restored prairie and exhibits similar sensitivity to drought as in remnant prairie. Furthermore, creating heterogeneity promotes forb productivity and enhances restored prairie resilience to drought. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is a necessary element of soil fertility and a limiting nutrient in tallgrass prairie but grazers like bison and cattle can also recycle N. Bison and cattle impact the nitrogen (N) cycle by digesting forage that is consumed, and recycled back to the soil in a more available forms stimulating soil microbial N cycling activities. Yet we do not know how both grazers comparatively affect N cycling in tallgrass prairie. Thus, we investigated if bison and cattle had similar impacts on N cycling in annually burned tallgrass prairie relative to ungrazed conditions over a 3-year period (2020–2022) at the Konza Prairie Biological Station. We examined: soil pH, soil water content, mineralized N, nitrification potential, denitrification potential and extracellular enzyme assays. Interannual variability in precipitation controlled soil water and N cycling microbial activities but grazing effects had a stronger influence on N cycling. We found significant differences and increased soil pH, nitrification and denitrification potential and less N limitation in bison vs cattle grazed soils where bison grazed soils exhibited faster N cycling. Differences between the grazers may be attributed to the different management of bison and cattle as both can impact N cycling. Overall, these data provide some evidence that bison and cattle affect N cycling differently at this study site, and improve the ecological understanding of grazer impacts on N cycling dynamics within the tallgrass prairie ecosystem. [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]

Tallgrass prairie is among the most threatened ecosystems but is often fragmented and surrounded by human‐modified landscapes. Small mammals are integral components of tallgrass prairies. However, little is known about how landscape composition, configuration, and management impact small mammals in tallgrass prairies.We conducted a systematic literature review to identify species‐specific and community associations with three broad topics: landscape composition, landscape configuration, and management practices.We identified 61 studies that assessed our variables of interest. We categorised the location, species assessed, variables monitored, and results by species and for the community.The majority of studies (64%) were conducted in two states, Illinois and Kansas. Deer mice (Peromyscus maniculatus), prairie voles (Microtus ochrogaster), and white‐footed mice (Peromyscus leucopus) showed specific associations with landscape variables, with deer mice preferring bare ground and recently burned plots, and prairie voles preferring thatch and negatively associated with prescribed fire. White‐footed mice were frequently associated with wooded areas.Small mammal biodiversity was positively associated with patchy habitats containing greater diversity in vegetative composition and management regime. Management and land composition were both relatively well studied for several species; habitat configuration was understudied.We identified significant gaps in our understanding of small mammal landscape ecology in tallgrass prairies. With tallgrass prairie restoration a growing trend in this region, a greater understanding of drivers of small mammal populations will be crucial to successful restoration efforts. Future research should focus on understudied areas and species, and examine how habitat heterogeneity impacts small mammal biodiversity. [ABSTRACT FROM AUTHOR]

Shrub encroachment is one of the primary threats to mesic grasslands around the world. This dramatic shift in plant cover has the potential to alter ecosystem‐scale water budgets and responses to novel rainfall regimes. Understanding divergent water‐use strategies among encroaching shrubs and the grasses they replace is critical for predicting shifts in ecosystem‐scale water dynamics as a result of shrub encroachment, particularly if drought events become more frequent and/or severe in the future.In this study, we assessed how water‐use traits of a rapidly encroaching clonal shrub (Cornus drummondii) and a dominant C4 grass (Andropogon gerardii) impact responses to changes in water availability in tallgrass prairie. We assessed intra‐annual change in depth of water uptake, turgor loss point and stomatal regulation in each species. Sampling took place at Konza Prairie Biological Station (northeastern KS, USA) during the 2021 and 2022 growing seasons.Cornus drummondii shifted from shallow to deep soil water sources across the 2021 and 2022 growing seasons. This plasticity in depth of water uptake facilitated a 'wasteful' water‐use strategy in C. drummondii, where stomatal conductance and transpiration rates continued to increase even when no further gain in photosynthetic rate occurred. A. gerardii photosynthetic rates and stomatal conductance were more variable through time and were more responsive to changes in leaf water potential than C. drummondii. However, intra‐annual adjustment of turgor loss point was more pronounced in C. drummondii (ΔπTLP = −0.48 MPa ± 0.15 SD) than in A. gerardii (ΔπTLP = −0.29 MPa ± 0.19 SD).Synthesis. These results suggest that C. drummondii is highly resilient to changes in water availability in surface soils and will likely remain unaffected by future droughts unless they are severe enough to reduce the availability of deep soil water. Given that clonal shrubs are key invaders of grasslands world‐wide, increased leaf‐level water loss is expected to accelerate ecosystem‐level drying as clonal shrub encroachment proceeds in mesic grasslands. [ABSTRACT FROM AUTHOR]

Animals must track resources over relatively fine spatial and temporal scales, particularly in disturbance‐mediated systems like grasslands. Grassland birds respond to habitat heterogeneity by dispersing among sites within and between years, yet we know little about how they make post‐dispersal settlement decisions. Many methods exist to quantify the resource selection of mobile taxa, but the habitat data used in these models are frequently not collected at the same location or time that individuals were present. This spatiotemporal misalignment may lead to incorrect interpretations and adverse conservation outcomes, particularly in dynamic systems. To investigate the extent to which spatially and temporally dynamic vegetation conditions and topography drive grassland bird settlement decisions, we integrated multiple data sources from our study site to predict slope, vegetation height, and multiple metrics of vegetation cover at any point in space and time within the temporal and spatial scope of our study. We paired these predictions with avian mark‐resight data for 8 years at the Konza Prairie Biological Station in NE Kansas to evaluate territory selection for Grasshopper Sparrows (Ammodramus savannarum), Dickcissels (Spiza americana), and Eastern Meadowlarks (Sturnella magna). Each species selected different types and amounts of herbaceous vegetation cover, but all three species preferred relatively flat areas with less than 6% shrub cover and less than 1% tree cover. We evaluated several scenarios of woody vegetation removal and found that, with a targeted approach, the simulated removal of just one isolated tree in the uplands created up to 14 ha of grassland bird habitat. This study supports growing evidence that small amounts of woody encroachment can fragment landscapes, augmenting conservation threats to grassland systems. Conversely, these results demonstrate that drastic increases in bird habitat area could be achieved through relatively efficient management interventions. The results and approaches reported pave the way for more efficient conservation efforts in grasslands and other systems through spatiotemporal alignment of habitat with animal behaviors and simulated impacts of management interventions. [ABSTRACT FROM AUTHOR]

We investigated the plant species diversity, community assemblage, net primary production and soil carbon sequestration in an old field (OF), which went through passive restoration with spontaneous secondary succession, and a restored prairie (RP), which went through active restoration with reintroduction of native plants. The passive restoration in OF did not attain as high diversity and richness of plant species as did in RP. Our NMS (nonmetric multidimensional scaling) ordination revealed a clear divergence of RP and OF vegetation with dissimilar species compositions to separate paths of succession trajectory. The spontaneous succession in OF has made very little progress due to a potentially strong inhibitory priority effect of invasive plants. The belowground biomass in the high‐diversity RP was 1.3× higher than the low‐diversity OF. We found no measurable difference in soil C sequestration between RP and OF at this time, possibly due to edaphic factors, lack of legumes, or both. Human intervention is needed for restoring abandoned croplands—highly disturbed, fragmented and isolated habitats—to tallgrass prairies in the U.S. Midwest. Our recommended intervention includes but is not limited to removal of exotic plants, further augmentation of native plants (particularly legumes), and periodic burns. [ABSTRACT FROM AUTHOR]

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One important goal in prairie restorations is to have forb species that flower throughout the growing season (i.e. a range of flowering phenology). We conducted an experiment to test whether three early‐ and three late‐flowering forb species are seed or recruitment limited, if mowing can influence this limitation, and if forb additions will impact overall plant diversity in ongoing restorations.The experiment was conducted at two restorations in northern Iowa and one in southern Minnesota, USA, using a split‐plot design. Whole plots tested the effect of mowing, and subplots tested the effect of target forbs being added as seeds or small transplants at varied richness levels (0, 1, 3 or 6 species added). Control plots received no forb additions. To compare establishment success, we estimated target species biomass, survivorship and per cent cover of target species across 3 years (2019–2021). Plant diversity measures (richness, inverse Simpson's diversity) were estimated within subplots to determine whether forb additions impacted diversity.Biomass and relative abundance of early‐ and late‐flowering forbs was much higher when they were added as transplants than when added as seeds. Survivorship across life stages indicated that the transition from seed to adult had lower survivorship probabilities (0.6%) than the transition from transplant (juvenile) to adult (75%).Plant diversity of these ongoing restorations was higher when target forbs were added compared to controls. Diversity was higher when target forbs were transplanted (8.4 species/plot and inverse Simpson's diversity of 4.0) than when they were seeded (7.6 species/plot and inverse Simpson's diversity of 3.8). Early‐flowering treatments increased diversity more than late‐flowering treatments, and diversity increased linearly with the number of forb species added.Mowed and unmowed plots had similar levels of target species establishment and plant diversity.Synthesis and applications: Our results indicate that early‐ and late‐flowering species are more recruitment limited than seed limited and that diversity is lower when these species are missing in restorations. We recommend transplanting a rich mix of early‐ and late‐flowering forbs to increase their biomass and ensure that flowering occurs throughout the growing season. [ABSTRACT FROM AUTHOR]

Root hemiparasites infiltrate the vascular tissue of host roots to acquire water and nutrients, which often reduces host growth. Hemiparasites are postulated to be keystone species in grassland communities if they suppress dominant species and increase plant community biodiversity, and ecosystem engineers if they increase nutrient accessibility for surrounding species. We examined keystone effects by evaluating species richness and evenness in 1 m2 plots in a recent prairie restoration where Castilleja sessiliflora was naturally present or absent, and in a longer-established prairie restoration with or without Pedicularis canadensis. We examined ecosystem engineer effects by determining nitrate and phosphate concentrations under, 25 cm from, and 50 cm from hemiparasites, and in the center of hemiparasite-free plots. On the C. sessiliflora site, plots with the hemiparasites had higher species richness due to more forbs and higher floristic quality, consistent with the keystone species hypothesis. Soil phosphate levels were also greater in plots with C. sessiliflora present, consistent with the hypothesis of ecosystem engineering by this hemiparasite. In contrast, plots with/without P. canadensis showed no associations of any community metrics with the hemiparasite, and no correspondence between the presence of hemiparasites and soil nutrients. Although hemiparasites can increase grassland community heterogeneity, the effect is not universal, and the direction and strength of effects likely depends on local conditions. [ABSTRACT FROM AUTHOR]

Climate change has initiated movement of both native and non‐native (exotic) species across the landscape. Exotic species are hypothesized to establish from seed more readily than comparable native species. We tested the hypothesis that seed limitation is more important for exotic species than native grassland species. We compared seed limitation and invasion resistance over three growing seasons between 18 native and 18 exotic species, grown in both monocultures and mixtures in a field experiment. Half of the plots received a seed mix of the contrasting treatment (i.e., exotic species were seeded into native plots, and native species were seeded into exotic plots), and half served as controls. We found that (1) establishment in this perennial grassland is seed limited, (2) establishment from seed is greater in exotic than native species, and (3) community resistance to seedling establishment was positively related to diversity of extant species, but only in native communities. Native‐exotic species diversity and composition differences did not converge over time. Our results imply that native to exotic transformations occur when diversity declines in native vegetation and exotic seeds arrive from adjacent sites, suggesting that managing for high diversity will reduce transformations to exotic dominance. [ABSTRACT FROM AUTHOR]

Experiments often find that net primary productivity (NPP) increases with species richness when native species are considered. However, relationships may be altered by exotic (non‐native) species, which are hypothesized to reduce richness but increase productivity (i.e., 'invasion‐diversity‐productivity paradox'). We compared richness‐NPP relationships using a comparison of exotic versus native‐dominated sites across the central USA, and two experiments under common environments. Aboveground NPP was measured using peak biomass clipping in all three studies, and belowground NPP was measured in one study with root ingrowth cores using root‐free soil. In all studies, there was a significantly positive relationship between NPP and richness across native species‐dominated sites and plots, but no relationship across exotic‐dominated ones. These results indicate that relationships between NPP and richness depend on whether native or exotic species are dominant, and that exotic species are 'breaking the rules', altering richness‐productivity and richness‐C stock relationships after invasion. [ABSTRACT FROM AUTHOR]

Plant-associated microbes, specifically fungal endophytes, augment the ability of many grasses to adapt to extreme environmental conditions. Tripsacum dactyloides (Eastern gamagrass) is a perennial, drought-tolerant grass native to the tallgrass prairies of the central USA. The extent to which the microbiome of T. dactyloides contributes to its drought tolerance is unknown. Ninety-seven genotypes of T. dactyloides were collected from native populations across an east–west precipitation gradient in Kansas, Oklahoma and Texas, and then grown together in a common garden for over 20 years. Root and leaf samples were visually examined for fungal density. Because fungal endophytes confer drought-tolerant capabilities to their host plants, we expected to find higher densities of fungal endophytes in plants from western, drier regions, compared to plants from eastern, wetter regions. Results confirmed a negative correlation between endophyte densities in roots and precipitation at the genotype's original location (r  = −0.21 P  = 0.04). Our analyses reveal that the host genotype's origin along the precipitation gradient predicts the absolute abundance of symbionts in the root, but not the relative abundances of particular organisms or the overall community composition. Overall, these results demonstrate that genetic variation for plant–microbe interactions can reflect historical environment, and reinforce the importance of considering plant genotype in conservation and restoration work in tallgrass prairie ecosystems. [ABSTRACT FROM AUTHOR]

Restoration of grassland ecosystems is essential for mitigating global losses of biodiversity and is typically initiated to foster persistent, long‐term increases in biodiversity. Yet, evaluating long‐term impacts of restoration on biodiversity is rare, especially across sites restored using consistent methods. Evaluation of restoration outcomes is particularly important for increasing predictive capacity in restoration ecology to determine the level of restoration effort that is required to achieve both short‐ and long‐term restoration goals. We conducted a multisite study that explicitly compared the impacts of no intervention ("passive" or "natural" recovery), low intervention (seeding native plants), and moderate intervention (seeding native plants and using fire management) at 32 restored prairies differing in the age of restoration (3–23 years). Grasslands with natural recovery have equivalent native plant species richness compared to sites with low and moderate restoration intervention, however, they have significantly lower‐quality vegetation, as measured by Floristic Quality (mean C). We found that managing restored prairies with fire maintains native plant richness over time and is correlated with higher vegetation quality and presence of seeded species. Seed mixes with a high mean C score are positively correlated with plant community quality. However, seed mixes with more species are negatively correlated with the proportion of seeded species present. We found that while the degree of restoration intervention has no effect on the number of native plant species, greater levels of assisted recovery are required to produce restored prairies that resemble high‐quality remnant vegetation and, especially, to maintain these successes over the long term. [ABSTRACT FROM AUTHOR]

Restoring habitat with diverse flowering plants is an important tool for pollinator conservation. Pollinator‐friendly seed mixes used in restorations vary in seeding density and forb‐to‐grass seed ratio, but few studies have examined the potential differences in nectar quantity and quality produced among mixes. We quantified the nectar rewards of Penstemon digitalis in plots planted with seed mixes of varying seeding density and forb‐to‐grass ratios by measuring nectar volume and sugar concentration. We found that nectar sugar concentration of P. digitalis was greatest at low seeding density, suggesting pollinator support in low‐cost seed mixes. Understanding how seed mix design parameters impact pollinator food resources is a key next step in habitat restoration for pollinator conservation. [ABSTRACT FROM AUTHOR]

Summary: Evolutionary history plays a key role driving patterns of trait variation across plant species. For scaling and modeling purposes, grass species are typically organized into C3 vs C4 plant functional types (PFTs). Plant functional type groupings may obscure important functional differences among species. Rather, grouping grasses by evolutionary lineage may better represent grass functional diversity.We measured 11 structural and physiological traits in situ from 75 grass species within the North American tallgrass prairie. We tested whether traits differed significantly among photosynthetic pathways or lineages (tribe) in annual and perennial grass species.Critically, we found evidence that grass traits varied among lineages, including independent origins of C4 photosynthesis. Using a rigorous model selection approach, tribe was included in the top models for five of nine traits for perennial species. Tribes were separable in a multivariate and phylogenetically controlled analysis of traits, owing to coordination of important structural and ecophysiological characteristics.Our findings suggest grouping grass species by photosynthetic pathway overlooks variation in several functional traits, particularly for C4 species. These results indicate that further assessment of lineage‐based differences at other sites and across other grass species distributions may improve representation of C4 species in trait comparison analyses and modeling investigations. [ABSTRACT FROM AUTHOR]

Phylogenetic diversity (PD), the evolutionary history of the organisms comprising a community, is increasingly recognized as an important driver of ecosystem function. However, biodiversity–ecosystem function experiments have rarely included PD as an a priori treatment. Thus, PD's effects in existing experiments are often confounded by covarying differences in species richness and functional trait diversity (FD). Here we report an experimental demonstration of strong PD effects on grassland primary productivity that are independent of FD, which was separately manipulated, and species richness, which was planted uniformly high to mimic diverse natural grasslands. Partitioning diversity effects demonstrated that higher PD increased complementarity (niche partitioning and/or facilitation) but lowered selection effects (probability of sampling highly productive species). Specifically, for every 5% increase in PD, complementarity increased by 26% on average (±8% SE), while selection effects decreased more modestly (8 ± 16%). PD also shaped productivity through clade‐level effects on functional traits, that is, trait values associated with particular plant families. This clade effect was most pronounced in the Asteraceae (sunflower family), which, in tallgrass prairies, generally comprises tall, high‐biomass species with low phylogenetic distinctiveness. FD also reduced selection effects but did not alter complementarity. Our results show that PD, independent of richness and FD, mediates ecosystem function through contrasting effects on complementarity and selection. This adds to growing evidence that consideration of phylogenetic dimensions of biodiversity can advance ecological understanding and inform conservation and restoration. [ABSTRACT FROM AUTHOR]

Ecological restoration outcomes are highly variable, undermining efforts to recover biodiversity and ecosystem functions. One poorly understood source of variability is 'year effects'—interannual variation in environmental conditions during the first year of restoration that alter successional trajectories of plant communities.There have been few experimental tests disentangling planting years from other differences among restoration projects (e.g. edaphic conditions, restoration approach), particularly those resolving mechanisms for year effects such as planting‐year rainfall. Moreover, past year effect studies focused almost exclusively on species‐level consequences. Therefore, the extent to which year effects influence the traits of communities is unknown.To address these gaps and provide a mechanistic test of how precipitation contributes to year effects, we conducted an experiment where we manipulated rainfall (drought, average and high levels) during the first growing season, replicated across three establishment year treatments to disentangle the effects of precipitation from other drivers of year effects. In each establishment year, we seeded the same species mix to initiate grassland restoration. We then surveyed plant community compositions annually for 5 years to quantify trait responses of restored communities to planting year rainfall.We found that variation in planting‐year precipitation altered community assembly trajectories by influencing community‐weighted mean (CWM) trait composition, and these effects persisted for at least 5 years. Over time, CWM specific leaf area and CWM seed mass decreased and CWM plant height increased. The effect of age on CWM plant height was stronger in plots that received mean and high watering treatments compared to drought treatments. This effect was also observed for CWM seed mass, albeit weaker.We also found some evidence for planting year effects unrelated to planting‐year rainfall for the three CWM traits, illustrating how interannually varying environmental conditions besides rainfall can generate persistent year effect on plant communities through their traits.Synthesis and applications. Our results provide evidence for planting year rainfall interacting with community assembly to alter the functional trait composition of restored grasslands. This suggests that interannual variation in rainfall during establishment is an important source of divergent biodiversity and functional outcomes in restored grasslands. [ABSTRACT FROM AUTHOR]

Grassland butterflies are undergoing worldwide population decline due to habitat loss and degradation. Rangelands in the Southern Great Plains can provide a habitat for grassland butterflies depending on management practices. Patch‐burn grazing is a management regime that involves burning portions of grazed pastures at different times. The combination of rotational fire and grazing creates a shifting mosaic of recently burned to older burned areas and lightly grazed to heavily grazed areas. However, the impact of fire and grazing on butterfly communities is complex and the effects of different management regimes on butterfly communities are not clear.We investigated the impact of time since prescribed fire and season of fire on butterfly communities in eight cattle‐grazed pastures, each with three burn units, in northern Oklahoma. Treatments included units burned in spring 2018, summer 2018, summer 2019, and spring 2020, with three replicates of each for a total of 12 burn units. Surveys were conducted three times per year in each burn unit in 2019 and 2020 using two standardised Pollard transects.A total of 909 butterflies and 35 species were observed. Species diversity varied by time since fire and season of fire, with spring‐burned sites having the lowest species diversity and summer‐burned sites having the highest. Dominant vegetation cover and blooming forb presence differed with time since fire and season of fire.Patch‐burn grazing creates a mosaic of successional vegetation stages which can benefit different butterfly species and support the overall community. Some species such as Cercyonis pegala, Danaus plexippus, and Atrytone arogos may benefit from fire every 1–2 years, while other species such as Echinargus isola may need longer times between fire treatments. Patch‐burn grazing regimes can support butterfly communities with species that need different fire return intervals by providing a mosaic of areas with different times since fire and associated grazing intensities. [ABSTRACT FROM AUTHOR]

Invasive species are suspected to be major contributors to biodiversity declines worldwide. Counterintuitively, however, invasive species effects are likely scale dependent and are hypothesized to be positively related to biodiversity at large spatial scales. Past studies investigating the effect of invasion on biodiversity have been mostly conducted at small scales (<100 m2) that cannot represent large dynamic landscapes by design. Therefore, replicated experimental evidence supporting a negative effect of invasive plants on biodiversity is lacking across many landscape types, including large grasslands.We collected data across eight large (333–809 ha) grassland landscapes managed with pyric herbivory—that is the recoupling of fire and grazing—to test how an invasive legume Lespedeza cuneata affected plant and bird communities at spatial grains ranging from 0.1 m2 to >3,000,000 m2.Lespedeza cuneata invasion effects on grassland plant diversity and composition changed with scale, being negative at small spatial grains (0.1 m2) and neutral or positive at large spatial grains (>3,000,000 m2).Lespedeza cuneata abundance did not significantly affect bird diversity at any spatial grain measured.Lespedeza cuneata may negatively affect biodiversity if abundances are greater than those observed in this study. However, previous research suggests that Lespedeza cuneata may not be capable of exceeding 20% canopy cover across large landscapes (>400 ha). Control and eradication strategies can be costly and are fraught with risk. If data do not clearly support a negative Lespedeza cuneata abundance–biodiversity relationship, and if invasion is spatially limited across large landscapes, ongoing control and eradication efforts may be unwarranted and ineffective.Synthesis and applications: Invasive species effects gleaned from small‐scale studies may not reliably predict their effects at larger scales. Although we recognize the importance of small‐scale studies in potentially isolating individual mechanisms, management strategies based solely on results from small‐scale studies of invasion are unlikely to increase or conserve biodiversity across large landscapes. Rather, processes that generate landscape heterogeneity—like pyric herbivory—are probably more important for promoting biodiversity across all scales. Scale is a central problem in ecology, and defining scale in management objectives is essential for effective biodiversity conservation. [ABSTRACT FROM AUTHOR]

Premise: Reproductive fitness in plants is often determined by the quantity and quality of pollen transferred by pollinators. However, many fitness studies measure only female fitness or rely on proxies for male fitness. Here we assessed how five bee taxon groups affect male fitness in a prairie plant by quantifying pollen removal, visitation, and siring success using paternity assignments and a unique pollinator visitation experiment. Methods: In Echinacea angustifolia, we measured per‐visit pollen removal for each pollinator taxon and estimated the number of pollen grains needed for successful ovule fertilization. Additionally, we directly measured pollinator influence on siring by allowing only one bee taxon to visit each pollen‐donor plant, while open‐pollinated plants acted as unrestricted pollen recipients. We genotyped the resulting offspring, assigned paternity, and used aster statistical models to quantify siring success. Results: Siring success of pollen‐donor plants differed among the five pollinator groups. Nongrooming male bees were associated with increased siring success. Bees from all taxa removed most of the flowering head's pollen in one visit. However, coneflower‐specialist bee Andrena helianthiformis removed the most pollen per visit. Female fitness and proxy measures of male fitness, such as pollinator visitation and pollen removal, did not align with our direct quantifications of male fitness. Conclusions: Our results illustrate the need for more studies to directly quantify male fitness, and we caution against using male fitness proxy measures. In addition, conservation efforts that preserve a diverse pollinator community can benefit plants in fragmented landscapes. [ABSTRACT FROM AUTHOR]

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The soil microbiome response to global change drivers remains largely unclassified, with current studies showing contrasting results to several global change drivers. One such driver, drought, is increasing in severity and frequency due to climate change. Further, legacy effects, or long‐lasting impacts after drought has subsided, could have lasting impacts on the soil microbiome with important consequences for ecosystem functioning. Thus, our study aimed to understand how the soil microbiome responds after a severe, prolonged drought and whether legacy effects persist post‐drought. We measured soil microbial community response in a mesic grassland for two years after a four‐year experimental drought in a native tallgrass prairie. The experimental drought was imposed either (1) chronically by reducing each growing season rainfall event by 66% or (2) intensely by completely eliminating growing season rainfall until an ~45% reduction in mean annual rainfall (climatic average) was achieved. The bacterial community showed no legacies in the first season post‐drought in response to either chronic or intense drought but showed an increased abundance of Verrucomicrobia and decreased richness in both treatments in the second growing season after the drought treatments ended. In the first and second post‐drought growing seasons, we found small differences in beta diversity between the control and intense drought treatment for fungal communities but not for the chronic drought treatment. Further, we found that the two main phyla of fungi, Ascomycota and Basidiomycota, showed reduced relative abundance post‐drought in the intense drought treatment. Overall, few legacies in soil microbial communities persisted after a four‐year experimentally induced drought. However, our results show that the nature of the drought—chronic versus intense—can differentially impact fungal versus bacterial short‐term legacies. These results suggest that the soil microbiome is for the most part drought resistant (responds little during drought) and/or resilient (quickly recovers post‐drought, since a study at the same site found that drought impacted the bacterial community) in this mesic grassland. [ABSTRACT FROM AUTHOR]

The conversion of tallgrass prairie to agriculture has negatively affected provisioning of ecosystem services. Successful restoration of ecosystem services could depend on management decisions applied during revegetation projects. We examined the effects of three management decisions (seed mix design, planting time, and first‐year mowing) on targeted ecosystem services (erosion control, weed resistance, and pollinator resources). We tested three seed mixes of varying diversity and grass‐to‐forb seeding ratios: Economy mix (21 species, 3:1 grass:forb), Pollinator mix (38 species, 1:3), and Diversity mix (71 species, 1:1). We established plots at two planting times (dormant‐season and spring) with or without first‐year mowing. To assess ecosystem services, we measured stem density, canopy cover, and floral density and richness of sown species in the second year after planting. The Economy mix had the highest stem density and cover but lowest floral density and richness. The Pollinator mix had the lowest stem density and cover but highest floral density. The Diversity mix had comparable stem density and cover to the Economy mix and comparable floral density and richness to the Pollinator mix. Mowing accelerated native plant establishment in all seed mixes. Dormant‐season planting improved establishment of spring and fall forbs and favored cool‐season graminoids over warm‐season grasses. All three management decisions influenced ecosystem outcomes, and comparison to a previous study revealed these effects to be robust to variation in site and climatic conditions. We recommend a diverse, balanced seed mix design, first‐year mowing, and dormant‐season planting to improve multifunctionality of conservation projects. [ABSTRACT FROM AUTHOR]

Potential natural vegetation (PNV) remains an important concept in ecology by providing the basis for comparing current or future conditions and trends in vegetation with historic conditions. However, nonstationary climate and land‐use‐driven deviations from baseline conditions challenge the validity of PNV at local scales or for locations that are especially sensitive to environmental changes, such as ecotonal boundaries between ecosystem types. Our goal was to identify the historic boundaries between major grassland types in the Central Great Plains (CGP) of North America (tallgrass prairie and northern and southern mixed‐grass prairies) by examining the environmental drivers of primary production prior to, during, and after the decades‐long drought in the 1930s. We sought to explore differences among grassland types in resistance and resilience to drought, and to evaluate legacy effects resulting from spatially explicit processes during drought. We used county‐level cropland production data from 1926 to 1948 as a proxy for native grass production data. We regressed production data against a suite of models containing variables associated with climate, soils, land use, and drought indices prior to, during, and after drought. We used a dimension reduction approach to identify the locations of grassland boundaries in multivariate space defined by each model. Our results show that climate and soil factors predicted grass production prior to the drought, yet additional land use and erosion factors were needed to explain production in both drought and post‐drought. Grassland responses were most variable along ecotonal boundaries. Tallgrass prairie exhibited stronger production responses to precipitation and temperature compared with northern and southern mixed‐grass prairies during all periods. Tallgrass prairie counties along the ecotone with mixed‐grass prairies were more negatively impacted by soil loss from abandoned cropland during the drought compared with other tallgrass prairie counties. As the intensity and frequency of drought are predicted to increase, landscape‐scale variables and spatially explicit processes that govern novel production responses will need to be accounted for in explaining heterogeneity in grassland responses. [ABSTRACT FROM AUTHOR]

Birds exhibit an assortment of behavioral strategies to cope with variable environmental conditions during reproduction, including altering nest construction behaviors. In species building enclosed domed nests, the microclimate within nests is influenced not only by its structure and the surrounding vegetation but also by the orientation of the nest opening. Many grassland-dependent birds build dome-shaped nests with clear directionality of openings. We studied two species in northeastern Kansas, United States that typically orient their nests east to northeast in this region. However, in a drought year, both Grasshopper Sparrows ( Ammodramus savannarum ) and Eastern Meadowlarks ( Sturnella magna ) shifted orientations of their nests southward toward prevailing winds. We hypothesized that this shift reduced the deleterious effects of heat stress on parents and developing young by diminishing morning solar radiation and increasing cooling due to the prevailing southerly winds. To test this hypothesis, we measured temperature, humidity, and wind speed at pairs of unoccupied, field-collected sparrow and meadowlark nests, experimentally placed to face south or east (control) in a non-drought year. Nest orientation affected the daily microclimate patterns, with south-facing nests warming later in the day relative to east-facing nests. The temperature differences depended upon humidity, with south-facing nests being relatively cooler under more humid conditions. This work provides the first experimental evidence of the benefits of plasticity in nest construction under challenging thermoregulatory conditions and shows how ground-nesting birds may reduce thermoregulatory demands during incubation under climate variation.

Root hemiparasites infiltrate the vascular tissue of host roots to acquire water and nutrients, which often reduces host growth. Hemiparasites are postulated to be keystone species in grassland communities if they suppress dominant species and increase plant community biodiversity, and ecosystem engineers if they increase nutrient accessibility for surrounding species. We examined keystone effects by evaluating species richness and evenness in 1 m2 plots in a recent prairie restoration where Castilleja sessiliflora was naturally present or absent, and in a longer-established prairie restoration with or without Pedicularis canadensis. We examined ecosystem engineer effects by determining nitrate and phosphate concentrations under, 25 cm from, and 50 cm from hemiparasites, and in the center of hemiparasite-free plots. On the C. sessiliflora site, plots with the hemiparasites had higher species richness due to more forbs and higher floristic quality, consistent with the keystone species hypothesis. Soil phosphate levels were also greater in plots with C. sessiliflora present, consistent with the hypothesis of ecosystem engineering by this hemiparasite. In contrast, plots with/without P. canadensis showed no associations of any community metrics with the hemiparasite, and no correspondence between the presence of hemiparasites and soil nutrients. Although hemiparasites can increase grassland community heterogeneity, the effect is not universal, and the direction and strength of effects likely depends on local conditions.

Taxonomic resolution is a critical component of biodiversity assessments. In this case study, we examined a single taxon within a larger study of nematode diversity to evaluate the taxonomic resolution of different diversity assessment methods. The selected taxon was the microbial-feeding genus Plectus, a group considered to include multiple cosmopolitan species. The methods included a morphological evaluation by light microscopy, Sanger sequencing of PCR amplicons of COI and 18S gene regions, and 18S metabarcoding sequencing. The study sites were 15 remnant tallgrass prairie plots in eastern Nebraska. In the morphological analysis, we observed two basic morphotypes, a short-tailed form with a small amphid and a long-tailed form with a large amphid. Sanger sequencing of COI sorted Plectus diversity into six distinct clades. The largest two of these six clades keyed to P. parietinus and P. rhizophilus based on morphology. BLAST analysis with COI revealed no close matches in GenBank. Sanger sequencing of the 18S region did not differentiate the six clades. These results illustrate that the method of diversity assessment strongly influences estimates of biodiversity. An additional 95 Plectus specimens, from outside the remnant sites, added taxonomic breadth to the COI phylogenetic tree. There were no geographically widespread COI haplotypes and no evidence of cosmopolitan Plectus species.

Abstract The soil microbiome response to global change drivers remains largely unclassified, with current studies showing contrasting results to several global change drivers. One such driver, drought, is increasing in severity and frequency due to climate change. Further, legacy effects, or long‐lasting impacts after drought has subsided, could have lasting impacts on the soil microbiome with important consequences for ecosystem functioning. Thus, our study aimed to understand how the soil microbiome responds after a severe, prolonged drought and whether legacy effects persist post‐drought. We measured soil microbial community response in a mesic grassland for two years after a four‐year experimental drought in a native tallgrass prairie. The experimental drought was imposed either (1) chronically by reducing each growing season rainfall event by 66% or (2) intensely by completely eliminating growing season rainfall until an ~45% reduction in mean annual rainfall (climatic average) was achieved. The bacterial community showed no legacies in the first season post‐drought in response to either chronic or intense drought but showed an increased abundance of Verrucomicrobia and decreased richness in both treatments in the second growing season after the drought treatments ended. In the first and second post‐drought growing seasons, we found small differences in beta diversity between the control and intense drought treatment for fungal communities but not for the chronic drought treatment. Further, we found that the two main phyla of fungi, Ascomycota and Basidiomycota, showed reduced relative abundance post‐drought in the intense drought treatment. Overall, few legacies in soil microbial communities persisted after a four‐year experimentally induced drought. However, our results show that the nature of the drought—chronic versus intense—can differentially impact fungal versus bacterial short‐term legacies. These results suggest that the soil microbiome is for the most part drought resistant (responds little during drought) and/or resilient (quickly recovers post‐drought, since a study at the same site found that drought impacted the bacterial community) in this mesic grassland.

Many exogenous factors may influence demographic rates (i.e., births, deaths, immigration, emigration), particularly for migratory birds that must cope with variable weather and habitat throughout their range and annual cycle. In midcontinental grasslands, disturbance (e.g., fire and grazing) and precipitation influence variation in grassland structure and function, but we know little about when and why precipitation is associated with grassland species' vital rates. We related estimates of detection, survival, and emigration to a priori sets of precipitation metrics to test the putative alternative factors influencing movement and mortality in grasshopper sparrows (Ammodramus savannarum). This species is a migratory songbird that exhibits exceptionally high rates of within‐season and between‐season dispersal. Between 2013 and 2020, we captured and resighted grasshopper sparrows in northeastern Kansas, USA, compiling capture histories for 1,332 adult males. We tested predictions of climatic hypotheses explaining variation in survival and emigration throughout a grasshopper sparrow's annual cycle; both survival and emigration were associated with the El Niño‐Southern Oscillation precipitation index (ESPI). Survival was positively related with ESPI during winter, and temporary emigration was curvilinearly related to breeding season ESPI lagged 2 years, with the highest site fidelity associated with intermediate rainfall values. The relationship between rainfall and temporary emigration likely reflects the influence of weather over multiple years on vegetation structure with consequent effects on local demography. This study provides compelling support for the idea that grassland species respond to high interannual variability by adopting dispersal strategies unlike those of many well‐studied migrant birds. Furthermore, the results imply that the consequences of increasing climatic extremes may not be immediately apparent, with demographic consequences lasting for at least a few years. [ABSTRACT FROM AUTHOR]

Questions: A robust ecosystem requires a functionally heterogeneous community of organisms with ecological traits that permit broad resource partitioning. Understanding community diversity patterns can help investigate drivers of community assembly and assess restoration success. Do biodiversity patterns differ among grassland communities sown with different ecotypes of dominant species during restoration along a rainfall gradient in the tallgrass prairie of the central US Great Plains? Location: Four field sites across a rainfall gradient within the North American Great Plains: Colby, Kansas (39°23′17.8″N, 101°04′57.4″W), Hays, Kansas (38°51′13.2″N, 99°19′08.6″W), Manhattan, Kansas (39°08′22.3″N, 96°38′23.3″W), and Carbondale, Illinois (IL, 37°41′47.0″N, 89°14′19.2″W). Methods: We applied linear mixed models to assess the effect of dominant species ecotype, year, and location on grassland taxonomic, phylogenetic, and functional diversity. Results: The non‐local grass ecotype (compared to the local ecotype) promoted species richness. In contrast, the effect of the dominant species ecotype on phylogenetic or functional diversity was site‐specific over the 10‐year restoration. Richness decreased across the rainfall gradient from dry to moist sites, and the wettest site had the highest phylogenetic and functional diversity. Conclusions: Our results suggest that abiotic filtering by rainfall is a key assembly mechanism that could predict grassland changes in biodiversity in the early restoration phases. Given the community response across the tallgrass prairie, restoration practitioners should consider the impact of regional sources of dominant species used in restoration when biodiversity is a restoration goal. It is recommended for future grassland restoration to detect gaps and limitations in evolutionary and trait structure that will reveal which diversity components to evaluate. [ABSTRACT FROM AUTHOR]

Plant community assembly outcomes can be contingent upon establishment year (year effects) due to variations in the environment. Stochastic events such as interannual variability in climate, particularly in the first year of community assembly, contribute to unpredictable community outcomes over the short term, but less is known about whether year effects produce transient or persistent states on a decadal timescale. To test for short‐term (5‐year) and persistent (decadal) effects of establishment year climate on community assembly outcomes, we restored prairie in an agricultural field using the same methods in four different years (2010, 2012, 2014, and 2016) that captured a wide range of initial (planting) year climate conditions. Species composition was measured for 5 years in all four restored prairies and for 9 and 11 years in the two oldest restored prairies established under average precipitation and extreme drought conditions. The composition of the four assembled communities showed large and significant differences in the first year of restoration, followed by dynamic change over time along a similar trajectory due to a temporary flush of annual volunteer species. Sown perennial species eventually came to dominate all communities, but communities remained distinct from each other in year five. Precipitation in June and July of the establishment year explained short‐term coarse community metrics (i.e., species richness and grass/forb cover), with wet establishment years resulting in a higher cover of grasses and dry establishment years resulting in a higher cover of forbs in restored communities. Short‐term differences in community composition, species richness, and grass/forb cover in restorations established under average precipitation and drought conditions persisted for 9–11 years, with low interannual variability in the composition of each prairie over the long term, indicating persistently different states on a decadal timescale. Thus, year effects resulting from stochastic variation in climate can have decadal effects on community assembly outcomes. [ABSTRACT FROM AUTHOR]

Woody encroachment has impacted grassland ecohydrology worldwide, prompting management strategies aimed at woody vegetation removal to prevent or mitigate loss of water yield. We measured stream discharge following sustained cutting of riparian trees (2010–2020) in a native tallgrass prairie (northeastern Kansas, USA). Discharge has declined at this site since the 1980's despite a concurrent increase in precipitation. This decline has been previously attributed to increased transpiration of stream water by riparian vegetation. We used water stable isotopes (δ18O and δ2H) to determine whether riparian grasses, shrubs, and trees primarily used stream/groundwater or soil water. Additionally, we quantified the increase in riparian and non-riparian woody cover (1978–2020) and combined it with sap-flux data to estimate changes in transpirative water loss. Sustained cutting of riparian trees did not result in increased discharge. Rather than stream/groundwater, the largest proportion of water used by riparian trees (Quercus spp.) was deep soil water. Cornus drummondii (clonal woody shrub) used a higher proportion of stream water and had greater overall variability in water-use. Riparian shrub cover increased about 57% from 1978 to 2020. Over the same time period, shrub cover increased about 20% in areas outside the riparian zone, resulting in an estimated 25% increase in daily transpirative water loss. Although stream water use was less than 50% for all riparian zone species, the total increase in shrub cover on this watershed, coupled with higher transpiration rates of shrubs, suggests that these woody species—within and outside the riparian zone—are key contributors to observed declines in stream flow in this system. [ABSTRACT FROM AUTHOR]

In grassland ecosystems, it is well known that increasing plant species diversity can improve ecosystem functions (i.e., ecosystem responses), for example, by increasing productivity and reducing weed invasion. Diversity-Interactions models use species proportions and their interactions as predictors in a regression framework to assess biodiversity and ecosystem function relationships. However, it can be difficult to model numerous interactions if there are many species, and interactions may be temporally variable or dependent on spatial planting patterns. We developed a new Diversity-Interactions mixed model for jointly assessing many species interactions and within-plot species planting pattern over multiple years. We model pairwise interactions using a small number of fixed parameters that incorporate spatial effects and supplement this by including all pairwise interaction variables as random effects, each constrained to have the same variance within each year. The random effects are indexed by pairs of species within plots rather than a plot-level factor as is typical in mixed models, and capture remaining variation due to pairwise species interactions parsimoniously. We apply our novel methodology to three years of weed invasion data from a 16-species grassland experiment that manipulated plant species diversity and spatial planting pattern and test its statistical properties in a simulation study.Supplementary materials accompanying this paper appear online. [ABSTRACT FROM AUTHOR]

Relationships between biodiversity and ecosystem functioning depend on the processes structuring community assembly. However, predicting biodiversity‐ecosystem functioning (BEF) relationships based on community assembly remains challenging because assembly outcomes are often contingent on history and the consequences of history for ecosystem functions are poorly understood. In a grassland restoration experiment, we isolated the role of history for the relationships between plant biodiversity and multiple ecosystem functions by initiating assembly in three different years, while controlling for all other aspects of community assembly. We found that two aspects of assembly history—establishment year and succession—altered species and trait community trajectories, which in turn altered net primary productivity, decomposition rates, and floral resources. Moreover, history altered BEF relationships (which ranged from positive to negative), both within and across functions, by modifying the causal pathways linking species identity, traits, diversity, and ecosystem functions. Our results show that the interplay of deterministic succession and environmental stochasticity during establishment mediate historical contingencies that cause variation in biodiversity and ecosystem functions, even under otherwise identical assembly conditions. An explicit attention to history is needed to understand why biodiversity‐ecosystem function relationships vary in natural ecosystems: a critical question at the intersection of fundamental theory and applications to environmental change biology and ecosystem restoration. [ABSTRACT FROM AUTHOR]

Moonlight structures activity patterns of many nocturnal species. Bright moonlight often limits the activity of nocturnal prey, but dense vegetation weakens this effect. Using 8 years of live‐trapping data, we asked whether reintroduced megaherbivores (Bison bison) indirectly altered moonlight avoidance by small mammals in tallgrass prairies. In plots with bison, plants intercepted 20% less light, allowing more moonlight to reach ground level. During nights with no moonlight, Peromyscus maniculatus activity was similar in plots with and without bison. During nights with peak moonlight, P. maniculatus activity was four times greater in plots without bison compared to plots with bison. Conversely, Microtus ochrogaster activity was twice as great during full moons compared to new moons, but only in plots with bison. We also equipped a subset of traps with temperature sensors to estimate trap‐entry time. Although M. ochrogaster was more active on bright nights, most activity occurred before moonrise or after moonset, avoiding periods of bright moonlight. We conclude that megaherbivores play an unappreciated but important indirect role in tallgrass prairies by inducing behavioral shifts in other animal species. Because overlap in activity patterns can predict the likelihood of predator–prey encounters, such activity shifts have important implications for trophic interactions throughout restored prairie food webs. Additional work to understand interspecific and intraspecific variation in response to moonlight may improve efforts to forecast changes in community assembly due to restoration and land‐use change. [ABSTRACT FROM AUTHOR]

Non-native earthworm invasions can greatly alter plant communities directly through root consumption, or indirectly, including through effects on arbuscular mycorrhizal (AM) fungi. However, most invasive earthworm research evaluating effects on plants and AM fungi has been conducted in historically earthworm-free regions. We assessed effects of non-native earthworms on plants and AM fungal abundance in a tallgrass prairie in the U.S. Great Plains where native earthworms are present. Specifically, we evaluated if abundance of native, non-native, and total earthworms were associated with native and non-native plant cover, plant biomass and species richness, and relative abundances of intra- and extra-radical AM fungi. We identified 68 non-native (all Aporrectodea spp.) and 451 native (all Diplocardia spp.) earthworms. Abundances of neither native nor non-native earthworms were associated with percent total plant cover, plant biomass, plant species richness, or intra- or inter-radical AM fungal abundance. However, greater numbers of total earthworms were associated with reduced native plant cover and increased non-native plant cover. Our study suggests total abundance of earthworms, but not of non-native species, influences plant community composition in the tallgrass prairie we studied, and that such effects may be driven by mechanisms other than alteration of symbiotic mycorrhizal fungi. As evidenced by the relatively low proportion of non-native individuals, the earthworm invasion in our study area appears to be nascent, yet further effects on soil, microbes, plants, and ecosystem functions could occur if the invasion intensifies. Continued research in tallgrass prairies is crucial for monitoring current and future impacts of non-native earthworms on these endangered ecosystems. [ABSTRACT FROM AUTHOR]

While the relationship between genetic diversity and plant productivity has been established for many species, it is unclear whether environmental conditions and biotic associations alter the nature of the relationship. To address this, we investigated the interactive effects of genotypic diversity, drought and mycorrhizal association on plant productivity and plant traits. Our mesocosm study was set up at the Konza Prairie Biological Research Station, located in the south of Manhattan, Kansas. Andropogon gerardii , the focal species for our study, was planted in two levels of genotypic richness treatment: monoculture or three-genotype polyculture. A rainout shelter was constructed over half of the experimental area to impose a drought and Thiophanate-methyl fungicide was used to suppress arbuscular mycorrhizal fungi in selected pots within each genotypic richness and drought treatment. Genotypic richness and mycorrhizal association did not affect above-ground biomass of A. gerardii. Drought differentially affected the above-ground biomass, the number of flowers and bolts of A. gerardii genotypes, and the biomass and the functional traits also differed for monoculture versus polyculture. Our results suggest that drought and genotypic richness can have variable outcomes for different genotypes of a plant species. [ABSTRACT FROM AUTHOR]

Context: Prairies historically covered much of inland North America, and many species have adapted to the unique conditions found in prairie ecosystems. Less than 1% of prairies remain in Arkansas, with much historic prairie having been converted for urban and agricultural development, resulting in steep population declines for many prairie-associated species. Objectives: Because many reptile and amphibian species are difficult to detect, the current distributions and habitat requirements of prairie-associated herpetofauna in fragmented landscapes are poorly understood. Thus, we assessed the state of prairie-associated herpetofauna communities in intact prairie, as well in degraded and developed historic prairie throughout Western Arkansas. Methods: Using repeated field surveys, remote sensing data, and hierarchical community occupancy models, we examined the influence of vegetation conditions, land use, and landscape characteristics on an assemblage of nine species of prairie-associated herpetofauna. Results: Prairie mound density, representing prairie that has not been subject to intense anthropogenic disturbance, was the strongest positive predictor of occupancy by prairie-associated species. Historic prairie area also exhibited a positive relationship with occupancy for several species but not at the assemblage level. Current vegetation conditions did not strongly influence occupancy patterns. Conclusions: Our results suggest that long-term land use filters, rather than present site-level conditions, are the driving forces dictating current distributions of prairie-associate herpetofauna in Western Arkansas. Our findings provide insight into the present state of understudied populations in an increasingly fragmented region and present accessible tools for directing exploratory conservation and research efforts. [ABSTRACT FROM AUTHOR]

Abstract During the “decade on restoration,” we must understand how to reliably re‐establish native plant populations. When establishing populations through seed addition, practitioners often prioritize obtaining seed from locations geographically near the restoration site (i.e. “local seed sourcing”). They are assumed to be under similar environmental conditions to the restoration site and should establish more robust plant populations and preserve local biotic interactions better than seeds sourced from further away. However, this assumption remains virtually untested in realistic restoration settings and the importance of seed sourcing, relative to other factors such as seeding rate and management regimes, is unclear. To determine if seed sourcing decision impacts plant establishment, abundance and phenology, we developed a partnership between university‐researchers and a native seed producer that kept records on where their seed was sourced from and where it was planted. At each site, we recorded the abundance and phenological stage of five commonly used tallgrass prairie restoration species seeded at 24 sites undergoing restoration across Michigan. We considered two measures of seed source locality: geographic distance (seeds were sourced from locations 6–750 km away from their respective restoration sites) and environmental distance. We also obtained data on the seeding rate and post‐seeding management at each site. We found that no measure of seed source locality predicted the likelihood of plant establishment or abundance at restoration sites. However, sites sown with seed from further away, or from cooler and wetter climates, had a greater proportion of flowering individuals earlier in the season. Finally, sites with higher seeding rates had greater plant abundance, and post‐seeding management of the restoration site increased the likelihood a species would establish by 36%. Overall, these results support that seed sourcing decisions did not impact plant establishment or abundance in our system. However, using less‐local seed sources may alter flowering phenology. Our results suggest that tallgrass prairie restoration efforts should prioritize higher seeding rates, post‐seeding management, and might expand the region seed sources are considered “local”, though this may impact flowering phenology. Future research leveraging native seed producer records can help answer critical questions about restoration seed sourcing.

Questions: Are community-weighted plant functional traits related to the 'fast-slow' growth rate continuum different in seed mixes, restored and reference prairies? If so, then what are the traits that explain differences? Remnant prairies, which serve as reference sites for restorations, often have higher plant species diversity and a lower abundance of exotic (non-native) species than their restoration counterparts. They may differ in their community-weighted functional traits as well, although this is poorly studied. Location: Tallgrass prairies and abandoned fields in Iowa, United States, North America. Methods: We compared functional traits between remnant prairies, restoration seed mixes, and established restorations to determine if community-weighted trait values (Spi × traits) are comparable. Relative abundance was determined by previous sampling of 93 restorations, 48 seed mixes and five remnant sites. Functional traits, including leaf specific leaf area (SLA) and leaf dry matter content (LDMC), and plant height, were measured for 145 native and 39 non-native plant species, 663 plants, and 1326 leaves. Results: Seed mixes had greater community-weighted LDMC and lower SLA than remnants, perhaps because of human selection for high grass abundance in mixes. Established restorations had lower community-weighted LDMC and higher SLA than remnants and seed mixes, and restorations had a large exotic species component. The proportional abundance of exotics in restorations explained trait differences between remnants and restorations. After taking into account functional groupings, perennial exotic species, on average, had higher SLA and lower LDMC than perennial native species. Remnants and established restorations had similar plant heights, and height was not significantly different between native and exotic species. Conclusions: Our results indicate that seed mixes differ from remnants in a manner that was the opposite of the difference between seed mixes and established restorations. Although seed mixes favored species with slower growth rate trait compositions, the species that established in restored prairies tended to have traits associated with faster growth rates. We recommend developing prairie restoration seed mixes with a greater proportion of forbs, especially those with lower LDMC and higher SLA. [ABSTRACT FROM AUTHOR]

Phylogenetic and functional diversity are relevant for restoration planning, as they influence important ecosystem functions and services. However, it is unknown whether initial phylogenetic and functional diversity of restorations as planned and planted are maintained over time, that is, the extent to which diversity of the restoration planting is reflected in the diversity of the resulting plant community. Furthermore, in the tallgrass prairie, many restorations are planted from seed. Among‐species variation in emergence and establishment affects the transition from seed mixes to realized plant communities in these restorations. We evaluated emergence and early establishment of experimental communities in a biodiversity plot experiment designed to test how phylogenetic and functional diversity influence restoration outcomes. We planted the same experimental communities starting from both seeds and plugs to assess differences in establishment. Our results suggest that phylogenetically and functionally diverse species mixes tend to produce phylogenetically and functionally diverse restored plant communities. After 3 years, experimental communities generally maintained their phylogenetic and functional diversity from seed and plug mixes to established vegetation, despite declines in species richness. While plots planted from seeds had on average 1.3 fewer species than plots planted from plugs, phylogenetic and functional diversity did not significantly differ between the two. Furthermore, most species exhibited no significant differences in percent cover when planted from seeds or plugs. Seeds are generally more cost‐effective for restoration than plugs, and our results indicate these two establishment methods achieved similar biodiversity outcomes. [ABSTRACT FROM AUTHOR]

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Mature ewes were used in a 4-yr study to evaluate effects of intensive late-season sheep grazing on vigor of sericea lespedeza in native tallgrass prairie. Pastures (N = 8; 31 ± 3.6 ha) infested with sericea lespedeza (initial basal frequency = 1.4%) were assigned randomly to one of two treatments: early-season beef steer grazing (1.1 ha/steer; initial BW = 258 ± 34 kg) from April 15 to July 15 followed by no grazing for the rest of the year (control; STR) or steer grazing from April 15 to July 15 followed by intensive grazing by mature ewes (0.2 ha/ewe; SHP) from August 1 to October 1. Ewes (initial BW = 65 ± 3.1 kg) were assigned randomly to graze four of eight pastures; remaining pastures were not grazed from August 1 to October 1. Vegetation responses to treatment were measured along four permanent 100-m transects in each pasture. Herbivory on sericea lespedeza was monitored weekly in each pasture from July 21 to October 7. Herbivory on sericea lespedeza in SHP and STR after steer grazing and before sheep grazing was not different (P = 0.51). In contrast, sericea lespedeza herbivory following sheep grazing was greater (P < 0.01) in SHP than in STR. Herbivory of individual sericea plants was greater (P < 0.01) in SHP than in STR by the end of week 1 of the sheep-grazing period (10.6% vs. 0.5%); moreover, herbivory on sericea lespedeza steadily increased (P ≤ 0.01) such that 92.1% of sericea lespedeza plants were grazed in SHP compared to 1.4% in STR by week 8 of the sheep-grazing period. Whole-plant DM weight of sericea lespedeza at dormancy was less (P < 0.01) in SHP than in STR. Additionally, annual seed production by sericea lespedeza was less (P < 0.01) in SHP than in STR (114 vs. 864 seeds/plant). Pasture forage biomass was not different (P = 0.76) between SHP and STR after the steer-grazing period. Conversely, STR had more (P < 0.01) residual forage biomass than SHP at the end of the sheep-grazing period. Growth performance of beef steers grazing from April 15 to July 15 annually was not different (P ≥ 0.59) between treatments. Our results were interpreted to suggest that intensive late-season grazing by sheep decreased vigor of sericea lespedeza. Late-season sheep grazing decreased forage biomass by 904 kg DM/ha compared with late-season rest; however, residual biomass was adequate to prevent soil-moisture loss and erosion during the dormant season. [ABSTRACT FROM AUTHOR]

Snakes are an integral component of grassland ecosystems, though their occurrence in high densities is often discouraged because of their role as nest predators of declining grassland birds. The effects of habitat structure on snake communities in grasslands remains poorly understood, hindering management efforts. We used 3 years of data from a series of restored grasslands in southern Illinois to examine how habitat structure across 3 spatial scales (local, patch, landscape) affected snake relative abundance, diversity, and species‐specific occupancy in grasslands. We found that snake community metrics were strongly and positively related to an increase in woody plant cover at the local (within‐patch) scale. Snake relative abundance was also positively related to an increase in grass cover and a decrease in forb cover, though our occupancy results suggest that this was primarily driven by an increase in black kingsnakes (Lampropeltis nigra). At the patch scale, relative abundance and diversity of snakes were both positively related to the proportion of patch edge composed of roads. Habitat structure at the landscape scale had the smallest impact on snakes in this study, though the proportion of trees in the landscape was positively related to snake diversity. We suggest that managers and conservationists interested in manipulating snake abundance in grasslands focus on within‐patch vegetation structure and composition. Decreasing woody cover in grasslands, or increasing the ratio of forbs to grasses, may reduce the presence of snakes, while maintaining a woody component could encourage both snake abundance and diversity. [ABSTRACT FROM AUTHOR]