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Questions: Woody plant encroachment and invasive plants are two critical factors that negatively impact grass‐dominated ecosystems. While studies have extensively investigated these factors individually, research on how invasive species impact the susceptibility of grasslands to encroachment is less common. Using the aspen parkland, an endangered savannah‐type ecosystem, we asked how the presence of a widespread invasive grass in North America, smooth brome (Bromus inermis), impacted the growth and survival of one‐year old trembling aspen (Populus tremuloides) seedlings. We also asked if plant litter was a potential mechanism of inhibition (as has been proposed for brome litter) or coexistence (as has been hypothesized for aspen litter) in this system. Location: Alberta, Canada. Methods: We used a manipulative experimental approach to determine if the survival and subsequent growth of planted aspen seedlings was impacted by the presence of smooth brome and manipulation of litter amount and type. Results: We found that the presence of smooth brome reduced aspen survival by 57% compared to uninvaded habitats, likely mediated by reduced soil moisture, while litter manipulation had no effect on survival. For surviving seedlings, local context had complex impacts on growth; the addition of aspen litter to brome‐invaded communities increased seedling growth while aspen litter additions to native communities resulted in decreased growth. Conclusion: These results suggest that invasion by smooth brome will alter the dynamics of aspen establishment in this system, potentially leading to significant changes to this already endangered landscape. Though smooth brome may serve as a barrier to aspen establishment, accumulation of aspen litter from nearby stands to brome patches could lead to faster growth of seedlings in invaded areas at the edge of existing aspen stands. Our results also suggest more generally that the impact of invasive plants on the establishment of native woody plants can be dependent on litter inputs. [ABSTRACT FROM AUTHOR]

Ecosystems are spatially heterogenous in plant community composition and function. Shrub occurrence in grasslands is a visually striking example of this, and much research has been conducted to understand the functional implications of this pattern. Within savannah ecosystems, the presence of tree and shrub overstories can have significant impacts on the understory herbaceous community. The exact outcomes, however, are likely a function of the spatial arrangement and traits of the overstory species. Here we test whether there are functional linkages between the spatial patterning of a native shrub and the standing biomass, community composition, and overall nutrient cycling of a neighbouring grassland understory communities within the Aspen Parkland of central Alberta, Canada. In a paired grassland-shrub stand study, we found the native shrub, Elaeagnus commutata , has relatively few stand-level impacts on the composition and standing biomass of the ecosystem. One factor contributing to these limited effects may be the overdispersion of shrub stems at fine spatial scales, preventing areas of deep shade. When we looked across a shrub density gradient and incorporated shrub architecture into our analyses, we found these shrub traits had significant associations with species abundance and root biomass in the understory community. These results suggest that stem dispersion patterns, as well as local stand architecture, are influential in determining how shrubs may affect their herbaceous plant understory. Thus, it is important to incorporate shrub spatial and architectural traits when assessing shrub-understory interactions. [ABSTRACT FROM AUTHOR]

Questions: Small‐scale disturbances have been linked to the maintenance of plant diversity by promoting the coexistence of competitive and subdominant species. Many proposed mechanisms for this coexistence involve changes to the competitive dynamics or abiotic environment. However, another possible mechanism could be a change to seed bank dynamics, ultimately impacting the species available for re‐establishment. Here, we test: (a) how gap creation alters seed bank composition, seed inputs (i.e. seed rain) and outputs (i.e. seedling establishment and predation); and (b) whether these changes are due to total vegetation removal or an indirect consequence of the removal of particularly influential species or traits. Location: Rough fescue grassland in Alberta, Canada. Methods: To test the impact of gap creation on seed bank dynamics, we experimentally removed or kept the standing vegetation intact in a paired‐plot design. Within each pair, we combined field sampling with greenhouse emergence studies to measure seed bank composition throughout the growing season. To examine inputs, we added seed trays that collected seed rain, and to examine outputs, we measured germination in seed trays with or without seed predation. Results: Seed bank composition within gaps was significantly different from the original standing vegetation before removal and the seed bank where standing vegetation was kept intact. Gap creation increased net seed rain and seedling establishment but had no impact on seed predation. These effects of gap creation on seed outputs seem to be reliant on the total removal of vegetation, while seed inputs are also dependent on the community and trait composition of the standing vegetation. Conclusions: The creation of gaps through small‐scale disturbances leading to a distinct seed bank community is another possible explanation for why we see increased β‐diversity with disturbance. Further, these gaps seem to be more important than standing vegetation compositional changes for preserving the local seed bank and promoting coexistence. [ABSTRACT FROM AUTHOR]

Questions: Local size structure is frequently measured in plant populations and forested systems due to its association with population‐ and community‐level processes. In contrast, size structure is rarely examined within herbaceous communities despite evidence for size‐dependent processes in these systems, suggesting an important functional vegetation property may be widely overlooked. Here, we test whether vertical size structure (a) varies within a short‐stature grassland, (b) varies along a productivity gradient, and (c) enhances the understanding of a foundational ecological pattern: the diversity–productivity relationship. Location: Rough fescue grassland in Alberta, Canada. Methods: We measured the heights of individuals within 32 plots and used average height, the Gini coefficient, and a two‐parameter Weibull distribution to estimate community‐level vertical size structure. Linear mixed models were used to test whether size structure parameters were significantly associated with productivity, species richness, and species evenness. Results: Vertical size structure varied significantly among local communities, such that some were dominated by short individuals while others were comprised of a more equal number of taller individuals. With increasing productivity, there was an increase in average plant height and a decrease in height inequality. Decreased height inequality was associated with reduced species richness with evidence that loss was not size‐dependent. The inclusion of vertical size structure parameters increased the explained variance of the diversity–productivity relationship by ~35% while also increasing its parsimony. Conclusions: Even within a short‐stature grassland, there is substantial functional variation in vertical size structure. Productivity is strongly associated with size structure patterns, and the inclusion of height inequalities greatly enhances productivity–diversity relationships, likely because they are the product of assembly mechanisms such as size‐asymmetric competition, assemblage‐level thinning, and niche complementarity. Overall, this demonstrates that vertical size structure has been a missing mechanism in most community assembly theories and models and should be included in the future. [ABSTRACT FROM AUTHOR]

Grasslands are important for C storage and supporting livestock production, yet little is understood about how warming interacts with defoliation to alter soil C and N dynamics. We examined how increased growing season temperature and defoliation influenced soil microbial biomass C (MBC) and N (MBN), N mineral dynamics, and greenhouse gas (GHG) flux within the top 15 cm of mineral soil in a northern temperate grassland of Alberta, Canada. We performed a 2-yr experiment with open-top chambers to increase soil temperatures, with warming assessed alone and combined with a single intense midsummer defoliation treatment. Soil MBC and MBN did not respond to warming or defoliation (P > 0.05). Warming increased soil nitrification and net N mineralization in the spring of the second growing season (P < 0.05) but did not persist beyond May. Although warming and defoliation both increased CO2 efflux separately during the first growing season (P < 0.05), the combination of these treatments reduced CO2 efflux to levels similar to the controls, with no lasting effects into the following growing season. Warming also interacted with defoliation to increase N2O flux in the second growing season relative to plots under either warming or defoliation treatment, whereas CH4 uptake was not influenced by our treatments. These results show the complex but transient impacts of warming and defoliation on soil C and N, particularly GHGs. Further research is needed to understand how these northern temperate grasslands respond to long-term factorial experiments involving warming, defoliation, and variation in other factors (e.g., precipitation) [ABSTRACT FROM AUTHOR]

The passenger, driver, and opportunist models are conceptual models of the invasion process used to describe alternative invasion scenarios. In the passenger model, both the invasive species and native community respond independently to environmental changes. In the driver model, changes to the native community are driven by the invasive species, while in the opportunist model invasion occurs in response to changes in the native community. In any given invasion scenario, however, it is possible that the relationships between the invasive, the native community, and the environment correspond to some combination of these invasion models acting simultaneously. We study invasion by Poa pratensis in a grassland in Alberta, Canada. Poa pratensis is a non-native plant implicated with loss of plant diversity in the region. In a three year field experiment, we manipulate the environment though defoliation, water addition, and nitrogen addition, and measure responses of P. pratensis cover, and cover and richness of the native community. We use structural equation modelling to describe the relationships between the invasive, the native community, and the environmental changes, and then interpret these relationships using the three invasion models. We found that P. pratensis predominantly invaded via the driver model, with subsequent reductions in native plant cover, but not in species richness. Positive effects of the environmental changes on P. pratensis also aided its ability to drive native cover. As well, we found some involvement of the opportunist model, through a negative relationship between the native community and the invasive. As invasion mainly proceeded via the driver model, management actions to limit invasion should focus on efforts to control abundance of P. pratensis itself. [ABSTRACT FROM AUTHOR]

We examined how water and nitrogen addition and water–nitrogen interactions affect root and shoot competition intensity and competition–productivity relationships in a native rough fescue grassland in central Alberta, Canada. Water and nitrogen were added in a factorial design to plots and root exclusion tubes and netting were used to isolate root and shoot competition on two focal species ( Artemisia frigida and Chenopodium leptophyllum). Both water and nitrogen were limiting to plant growth, and focal plant survival rates increased with nitrogen but not water addition. Relative allocation to root biomass increased with water addition. Competition was almost entirely belowground, with focal plants larger when released from root but not shoot competition. There were no significant relationships between productivity and root, shoot, or total competition intensity, likely because in this system shoot biomass was too low to cause strong shoot competition and root biomass was above the levels at which root competition saturates. Water addition had few effects on the intensity of root competition suggesting that root competition intensity is invariant along soil moisture gradients. Contrary to general expectation, the strength of root competition increased with nitrogen addition demonstrating that the relationship between root competition intensity and nitrogen is more complex than a simple monotonic decline as nitrogen increases. Finally, there were few interactions between nitrogen and water affecting competition. Together these results indicate that the mechanisms of competition for water and nitrogen likely differ. [ABSTRACT FROM AUTHOR]

Summary A field experiment in a native grassland in Central Alberta, Canada, tested whether plants alter relative allocation to roots vs. shoots in response to below-ground competition, and whether the mass of a species’ root system accounts for interspecific differences in below-ground competitive response. 2 Seedlings of each of 10 native species were transplanted into the naturally occurring vegetation in the field at the start of the growing season. Root interactions between the target plants and their neighbours were manipulated through the use of PVC root exclusion tubes, with target plants grown with or without potential root interactions with their neighbours. Neighbour shoots were also tied back, forcing any target–neighbour interactions to be below ground. 3 Below-ground competition generally reduced plant growth, with its relative magnitude varying among species. 4 An allometric analysis indicated that competition below ground did not result in an increase in the relative biomass allocated to roots for any of the 10 target species. This is counter to the growth-balance hypothesis (and optimal foraging theory). Below-ground competition did increase root : shoot ratios, but this was due to reduced plant size (small plants have larger root : shoot ratios), rather than adaptive plasticity. 5 A species’ below-ground competitive ability was not related to its root system size. Although this finding is counter to commonly made assumptions, it is supported by other work demonstrating below-ground competition to be generally size-symmetric. 6 Despite the majority of plant–plant interactions in grasslands being below ground, our understanding of plant competition above ground is significantly more robust. Several wide-spread assumptions regarding below-ground competition are suspect, and more multispecies studies such as this are required to provide a fuller picture of how plants respond to, and compete for, soil resources. [ABSTRACT FROM AUTHOR]

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Invasive species can alter the structure and function of the communities they invade, as well as lead to biotic homogenization across their invasive range, thus affecting large‐scale diversity patterns. The mechanisms by which invasive species can lead to biotic homogenization are poorly understood. We argue that invasive species acting as strong, deterministic, and consistent filters within and across invaded communities are likely to cause biotic homogenization at multiple spatial scales. We studied Bromus inermis Leyss. invasion into eight grasslands covering most of the grassland and parkland natural regions of Alberta (western Canada). Specifically, we tested whether B. inermis (1) has a strong impact on species richness and composition, (2) consistently alters resources (nutrients, light, and soil moisture), imposing the same ecological filter to species establishment and persistence across sites, and hence (3) whether it leads to biotic homogenization within and across sites. We recorded plant cover and resources across native‐to‐invaded transition areas combining space‐for‐time substitutions with time‐series data analyses. Bromus inermis invasion was associated with rapid biotic homogenization of communities, within and among the eight grasslands. The sharp changes in species relative abundances following invasion was the initial driver of biotic homogenization, and species loss was delayed. Supporting the idea that biotic homogenization can occur when an invasive species presents a broad and consistent filter, resources modified by B. inermis invasion (particularly light and certain nutrients) were altered rather consistently within and across sites. The 50% reduction in light was likely the initial driver of biotic homogenization, and the increase in nutrient availability probably facilitates the displacement of species from the invaded areas and could lead to the establishment of self‐reinforcing dynamics. Overall, our results support the idea that invaders acting as strong, deterministic, and consistent ecological filters are likely to cause biotic homogenization of the communities they invade. [ABSTRACT FROM AUTHOR]

• We studied in-situ responses of GHGs to warming, altered precipitation and defoliation. • The temperate grassland was a source for CO 2 , N 2 O but a sink for CH 4 during the study. • GHG fluxes were decoupled from microbial biomass in the study. • Precipitation was the primary driver directly regulating CO 2 and N 2 O fluxes. Grasslands store large quantities of organic carbon (C) and thus can markedly impact global climate change if they are disturbed. How practices such as grazing intensity and climate change interact to affect soil greenhouse gas (GHG) fluxes within native northern temperate grasslands is unclear. We investigated the responses of three major GHGs (CO 2 , N 2 O and CH 4) to temperature (ambient vs. warmed), precipitation (ambient vs. reduced or increased) and defoliation (no defoliation vs. low- or high-intensity defoliation) treatments in a factorial experiment over a two-year period in a native northern temperate grassland in Alberta, Canada. Growing season mean soil gas fluxes across all treatments were 93.8 mg CO 2 m−2 h−1, −0.025 mg CH 4 m−2 h−1, and 0.003 mg N 2 O m−2 h−1, suggesting that this grassland was a source for CO 2 and N 2 O, but a sink for CH 4 over the study period. Structural equation modeling showed that increased precipitation, rather than warming or defoliation, was the primary driver directly regulating fluxes of CO 2 and N 2 O. While increased precipitation also increased microbial biomass C (MBC), the latter had little impact on GHG fluxes, suggesting that MBC was not a critical factor regulating the precipitation-GHG flux relationship. Warming and precipitation effects on CH 4 fluxes depended on the month of sampling during the growing season. Warming decreased CH 4 uptake but only in mid-July, while increased precipitation reduced CH 4 uptake across all sampling dates except early August, at which time increased precipitation increased CH 4 uptake. Our study shows that short-term soil GHG fluxes in this native grassland are relatively resilient to moderate warming and defoliation, but sensitive to altered precipitation. Increased precipitation was the main contributor to short-term GHG fluxes in this temperate grassland via changes in CO 2 efflux. [ABSTRACT FROM AUTHOR]

Background: Localized disturbances within grasslands alter biological properties and may shift species composition. For example, rare species in established communities may become dominant in successional communities if they exhibit traits well-suited to disturbance conditions. Although the idea that plant species exhibit different trait 'strategies' is well established, it is unclear how ecological selection for specific traits may change as a function of disturbance. Further, there is little data available testing whether disturbances select for single trait-characters within communities (homogenization), or allow multiple trait-types to persist (diversification). We investigated how (a) traits and (b) functional diversity of post-disturbance gap communities compared to those in adjacent undisturbed grasslands, and (c) if altered functional diversity resulted in the homogenization or diversification of functional traits., Methods: Here we emulate the impacts of an extreme disturbance in a native grassland site. We measured plant community composition of twelve paired 50 × 50 cm plots (24 total) in Alberta, Canada. Each pair consisted of one undisturbed plot and one which had all plants terminated 2 years prior. We used species abundances and a local trait database to calculate community weighted means for maximum height, specific leaf area, specific root length, leaf nitrogen percent, and root nitrogen percent. To test the impacts of disturbance on community functional traits, we calculated functional diversity measures and compared them between disturbed and undisturbed communities., Results: Within 2 years, species richness and evenness in disturbed communities had recovered and was equivalent to undisturbed communities. However, disturbed and undisturbed communities had distinct community compositions, resulting in lower functional divergence in disturbed plots. Further, disturbance was linked to increases in community-weighted mean trait values for resource-acquisitive traits, such as specific leaf area, and leaf and root nitrogen., Discussion: Disturbance had lasting effects on the functional traits and diversity of communities, despite traditional biodiversity measures such as richness and evenness recovering within 2 years. The trait space of gap communities shifted compared to undisturbed communities such that gap communities were dominated by traits enhancing resource uptake and growth rates. Overall, these results show that short-term disturbance fundamentally changes the functional character of early-successional communities, even if they superficially appear recovered., Competing Interests: The authors declare that they have no competing interests., (© 2022 Smith et al.)

A primary impediment to understanding how species diversity and anthropogenic disturbance are related is that both diversity and disturbance can depend on the scales at which they are sampled. While the scale dependence of diversity estimation has received substantial attention, the scale dependence of disturbance estimation has been essentially overlooked. Here, we break from conventional examination of the diversity-disturbance relationship by holding the area over which species richness is estimated constant and instead manipulating the area over which human disturbance is measured. In the boreal forest ecoregion of Alberta, Canada, we test the dependence of species richness on disturbance scale, the scale-dependence of the intermediate disturbance hypothesis, and the consistency of these patterns in native versus exotic species and among human disturbance types. We related field observed species richness in 1 ha surveys of 372 boreal vascular plant communities to remotely sensed measures of human disturbance extent at two survey scales: local (1 ha) and landscape (18 km2). Supporting the intermediate disturbance hypothesis, species richness-disturbance relationships were quadratic at both local and landscape scales of disturbance measurement. This suggests the shape of richness-disturbance relationships is independent of the scale at which disturbance is assessed, despite that local diversity is influenced by disturbance at different scales by different mechanisms, such as direct removal of individuals (local) or indirect alteration of propagule supply (landscape). By contrast, predictions of species richness did depend on scale of disturbance measurement: with high local disturbance richness was double that under high landscape disturbance.

Background: Niche theory predicts that human disturbance should influence the assembly of communities, favouring functionally homogeneous communities dominated by few but widespread generalists. The decline and loss of specialists leaves communities with species that are functionally more similar. Evenness of species occupancy declines, such that species become either widespread of rare. These patterns have often been observed, but it is unclear if they are a general result of human disturbance or specific to communities that are rich in species, in complex, spatially heterogeneous environments where the problem has often been investigated. We therefore tested whether human disturbance impacts dominance/evenness of species occupancy in communities, specialism/generalism of species, and functional biotic homogenization in the spatially relatively homogeneous, species poor boreal forest region of Alberta, Canada. We investigated 371 boreal vascular plant communities varying 0 - 100% in proportion of human land use., Results: Rank species occupancy curves revealed high species dominance regardless of disturbance: within any disturbance class a few species occupied nearly every site and most species were found in a low proportion of sites. However, species were more widespread and displayed more even occupancy in intermediately disturbed communities than among communities of either low or high disturbance. We defined specialists and generalists based on turnover in co-occupants and thereby assessed impacts of human disturbance on specialization of species and community homogenization. Generalists were not disproportionately found at higher disturbance sites, and did not occupy more sites. Communities with greater human disturbance were not more functionally homogeneous; they did not harbor communities with more generalists., Conclusions: We unexpectedly did not observe strong linkages between species specialism/generalism and disturbance, nor between community homogenization and disturbance. These results contrast previous findings in more species rich, complex or spatially heterogeneous systems and ecological models. We suggest that broad occupancy-based intercommunity patterns are insensitive to human land use extent in boreal vascular plants, perhaps because of ubiquity of generalists, low species richness, and history of natural disturbance. The poor sensitivity of these metrics to disturbance presents challenges for monitoring and managing impacts to biodiversity in this region.

Ungulates impact woody species' growth and abundance but little is understood about the comparative impacts of different ungulate species on forest expansion in savanna environments. Replacement of native herbivore guilds with livestock [i.e., beef cattle (Bos taurus)] has been hypothesized as a factor facilitating trembling aspen (Populus tremuloides Michx.) encroachment into grasslands of the Northern Great Plains. We used a controlled herbivory study in the Parklands of western Canada to compare the impact of native ungulates and cattle on aspen saplings. Native ungulate treatments included a mixed species guild and sequences of herbivory by different ungulates [bison (Bison bison subsp. bison), elk (Cervus elaphus) then deer (Odocoileus hemionus); or deer, elk, then bison]. Herbivory treatments were replicated in three pastures, within which sets of 40 marked aspen saplings (<1.8 m) were tracked along permanent transects at 2-week intervals, and compared to a non-grazed aspen stand. Stems were assessed for mortality and incremental damage (herbivory, leader breakage, stem abrasion and trampling). Final mortality was greater with exposure to any type of herbivore, but remained similar between ungulate treatments. However, among all treatments, the growth of aspen was highest with exposure only to cattle. Herbivory of aspen was attributed primarily to elk within the native ungulate treatments, with other forms of physical damage, and ultimately sapling mortality, associated with exposure to bison. Overall, these results indicate that native ungulates, specifically elk and bison, have more negative impacts on aspen saplings and provide evidence that native and domestic ungulates can have different functional effects on woody plant dynamics in savanna ecosystems.