Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

Direct and indirect effects of environmental factors,spatial constraints,and functional traits on shaping the plant diversity of montane forests

Understanding the relative importance of the factors driving the patterns of biodiversity is a key research topic in community ecology and biogeography. However, the main drivers of plant species diversity in montane forests are still not clear. In addition, most existing studies make no distinction between direct and indirect effects of environmental factors and spatial constraints on plant biodiversity. Using data from 107 montane forest plots in Sichuan Giant Panda habitat, China, we quantified the direct and indirect effects of abiotic environmental factors, spatial constraints, and plant functional traits on plant community diversity. Our results showed significant correlations between abiotic environmental factors and trees (r = .10, p value = .001), shrubs (r = .19, p value = .001), or overall plant diversity (r = .18, p value = .001) in montane forests. Spatial constraints also showed significant correlations with trees and shrubs. However, no significant correlations were found between functional traits and plant community diversity. Moreover, the diversity (richness and abundance) of shrubs, trees, and plant communities was directly affected by precipitation, latitude, and altitude. Mean annual temperature (MAT) had no direct effect on the richness of tree and plant communities. Further, MAT and precipitation indirectly affected plant communities via the tree canopy. The results revealed a stronger direct effect on montane plant diversity than indirect effect, suggesting that single‐species models may be adequate for forecasting the impacts of climate factors in these communities. The shifting of tree canopy coverage might be a potential indicator for trends of plant diversity under climate change.     

Neighbourhood association of Cortaderia selloana invasion,soil properties and plant community structure in Mediterranean coastal grasslands

Invasion by alien species is threatening the conservation of native plant communities and the integrity of ecosystems. To gain a better understanding of such impacts, many studies have examined the traits that make alien species successful invaders as well as the factors involved in community invasibility. However, it is necessary to link invader effects on community structure and on ecosystem processes in order to unravel the mechanisms of impact. Cortaderia selloana is a perennial grass native to South America that is invading abandoned agricultural lands close to coastal human settlements in Catalonia (NE Spain). In invaded pastures, we examined the association between C. selloana invasion, soil properties and vegetation structure changes in pastures, comparing the neighbourhood area of influence of C. selloana with areas far from C. selloana. Areas under the influence of C. selloana had lower total soil nitrogen values and higher C/N values than in areas far from C. selloana. Furthermore, the areas affected by C. selloana had lower species, family and life form richness and diversity, and less plant cover. In addition, C. selloana also increased the vertical vegetation structure and changed species composition (only 44% similarity between invaded and non-invaded areas). Our results point out that C. selloana has an effect on its neighbourhood leading to an increase in small-scale variability within invaded fields.     

Feral horses dung piles as potential invasion windows for alien plant species in natural grasslands

Small scale disturbances could act as patches that provide sites for the colonization of competitively inferior species, promoting the establishment of non-native species in some cases. We analyzed the vegetation associated with feral horse dung piles in montane pampas grasslands in Mid-East Argentina and described the changes following their abandonment, evaluating whether dung piles act as invasion windows, allowing the entrance of alien plant species. We estimated the portion of the study area directly covered by horse manure and dung height was used to estimate the time elapsed after the abandonment of each pile. Vegetation replacement on dung piles of different ages was assessed and compared with grassland controls using discriminant analysis. We used regression analysis to look for changes in vegetation cover, species richness, species diversity and evenness in response to height (age) of the dung piles, and principal component analyses (PCA) to identify groups of plants associated with different successional stages. We compared cover of alien plant species on dung piles with grassland controls using one-way ANOVA. On average, 2.5% of the study area was covered by horse dung. Total vegetation cover, species richness, diversity and evenness increased after the piles were abandoned. Characteristic plant groups were associated with initial, middle and last phases of the studied succession. Vegetation on the dung piles significantly differed from that in grassland controls and two species were consistently associated with dung piles: the invasive Red Star Thistle, Centaurea calcitrapa, and a native grazing-intolerant grass, Nassella clarazii. Non-native species cover was also higher in dung piles than in control plots. Dung piles cover a significant portion of grassland area in our study site, produce significant changes in the vegetation and are associated with some invasive alien plants that could eventually colonize more pristine areas in the vicinity. On the other hand, they might represent refuges for palatable species, since horses seem to avoid them for grazing.     

Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Relationship between native and exotic plant species at multiple savannah sites

We tested whether the recently proposed two‐part measure of degree of invasion (DI) of a community relating exotic proportion of cover to exotic proportion of richness can characterize patterns of plant invasions at multiple savannah sites in Southern Africa. Regression analysis was performed on transformed data to assess how this two‐part measure of DI compares to other metrics of community invasibility. The results indicate that at the plot level, the absolute cover of exotics was not significantly related to native cover for three sites out of four assessed (R² ≤ 0.17; P > 0.05). Also, at all four sites, no significant relationships were detected between native and exotic plant richness at both the 1‐m² and 400‐m² plot scales. By contrast, significant (P < 0.05) positive linear relationships were observed between exotic proportion of richness and exotic proportion of cover at all sites (R² was as high as 0.67 and 0.97 for two sites). Our results indicate that the new two‐part measure of DI is able to characterize patterns of plant invasions across plant communities in African savannahs.     

Arbuscular mycorrhizal inoculum potential: a mechanism promoting positive diversity�Cinvasibility relationships in mountain beech forests in New Zealand?

Mycorrhizal fungi are important symbionts for the majority of plant species, but their role in determining the susceptibility of habitat to plant invasion is poorly understood. Hieracium lepidulum is an arbuscular mycorrhizal herb, currently invading the understorey of ectomycorrhizal Nothofagus solandri var. cliffortioides (mountain beech) forest in New Zealand. Mountain beech is solely ectomycorrhizal, and other plant species within the understorey occur sporadically. Hieracium has been shown to establish preferentially in microsites with higher plant species richness at a scale of less than 1 m² within mountain beech forest, and we tested the hypothesis that more diverse microsites (<1 m²) are associated with higher levels of arbuscular mycorrhizal fungal (AMF) inoculum. We found low levels of AMF inoculum across all microsites, and over a third of samples contained no inoculum at all. Higher vascular-plant species richness (but not biomass) was associated with higher AMF spore densities in field soil, and greater AMF colonization of H. lepidulum seedlings in a bioassay. Absence of AMF inoculum from much of the soil and the positive association of inoculum potential with species richness provide a potential mechanism for the establishment of a positive diversity–invasibility relationship in the mountain beech forest.     

Effects of plant species richness and evenness on soil microbial community diversity and function

Understanding the links between plant diversity and soil communities is critical to disentangling the mechanisms by which plant communities modulate ecosystem function. Experimental plant communities varying in species richness, evenness, and density were established using a response surface design and soil community properties including bacterial and archaeal abundance, richness, and evenness were measured. The potential to perform a representative soil ecosystem function, oxidation of ammonium to nitrite, was measured via archaeal and bacterial amoA genes. Structural equation modeling was used to explore the direct and indirect effects of the plant community on soil diversity and potential function. Plant communities influenced archaea and bacteria via different pathways. Species richness and evenness had significant direct effects on soil microbial community structure, but the mechanisms driving these effects did not include either root biomass or the pools of carbon and nitrogen available to the soil microbial community. Species richness had direct positive effects on archaeal amoA prevalence, but only indirect impacts on bacterial communities through modulation of plant evenness. Increased plant evenness increased bacterial abundance which in turn increased bacterial amoA abundance. These results suggest that plant community evenness may have a strong impact on some aspects of soil ecosystem function. We show that a more even plant community increased bacterial abundance, which then increased the potential for bacterial nitrification. A more even plant community also increased total dissolved nitrogen in the soil, which decreased the potential for archaeal nitrification. The role of plant evenness in structuring the soil community suggests mechanisms including complementarity in root exudate profiles or root foraging patterns.     

Evenness–invasibility relationships differ between two extinction scenarios in tallgrass prairie

Experiments that have manipulated species richness with random draws of species from a larger species pool have usually found that invasibility declines as richness increases. These results have usually been attributed to niche complementarity, and interpreted to mean that communities will become less resistant to invaders as species go locally extinct. However, it is not clear how relevant these studies are to real‐world situations where species extinctions are non‐random, and where species diversity declines due to increased rarity (i.e. reduced evenness) without having local extinctions. We experimentally varied species richness from 1 to 4, and evenness from 0.44 to 0.97 with two different extinction scenarios in two‐year old plantings using seedling transplants in western Iowa. In both scenarios, evenness was varied by changing the level of dominance of the tall grass Andropogon gerardii. In one scenario, which simulated a loss of short species from Andropogon communities, we directly tested for complementarity in light capture due to having species in mixtures with dissimilar heights. We contrasted this scenario with a second set of mixtures that contained all tall species. In both cases, we controlled for factors such as rooting depth and planting density. Mean invader biomass was higher in monocultures (5.4 g m^?2 week^?1) than in 4‐species mixtures (3.2 g m^?2 week^?1). Reduced evenness did not affect invader biomass in mixtures with dissimilar heights. However, the amount of invader biomass decreased by 60% as evenness increased across mixtures with all tall species. This difference was most pronounced early in the growing season when high evenness plots had greater light capture than low evenness plots. These results suggest that the effect of reduced species diversity on invasibility are 1) not related to complementarity through height dissimilarity, and 2) variable depending on the phenological traits of the species that are becoming rare or going locally extinct.     

Spatial heterogeneity of soil nutrients and plant species in herb-dominated communities of contrasting land use

Recent interest in spatial pattern in terrestrial ecosystems has come from an awareness of the intimate relationship between spatial heterogeneity of soil resources and maintenance of plant species diversity. Soil and vegetation can vary spatially in response to several state factors of the system. In this study, we examined fine-scale spatial variability of soil nutrients and vascular plant species in contrasting herb-dominated communities (a pasture and an old field) to determine degree of spatial dependence among soil variables and plant community characteristics within these communities by sampling at 1-m intervals. Each site was divided into 25 1-m² plots. Mineral soil was sampled (2-cm diameter, 5-cm depth) from each of four 0.25-m² quarters and combined into a single composite sample per plot. Soil organic matter was measured as loss-on-ignition. Extractable NH₄ and NO₃ were determined before and after laboratory incubation (28 days at 27°C) to determine potential net N mineralization and nitrification. Cations were analyzed using inductively coupled plasma emission spectrometry. Vegetation was assessed using estimated percent cover. Most soil and plant variables exhibited sharp contrasts between pasture and old-field sites, with the old field having significantly higher net N mineralization/nitrification, pH, Ca, Mg, Al, plant cover, and species diversity, richness, and evenness. Multiple regressions revealed that all plant variables (species diversity, richness, evenness, and cover) were significantly related to soil characteristics (available nitrogen, organic matter, moisture, pH, Ca, and Mg) in the pasture; in the old field only cover was significantly related to soil characteristics (organic matter and moisture). Both sites contrasted sharply with respect to spatial pattern of soil variables, with the old field exhibiting a higher degree of spatial dependence. These results demonstrate that land-use practices can exert profound influence on spatial heterogeneity of both soil properties and vegetation in herb-dominated communities.     

Effects of belowground resource use comlementarity on invasion of constructed grassland plant communities

At two field sites that differed in fertility, we investigated how species richness, functional group diversity, and species composition of constructed plant communities influenced invasion. Grassland communities were constructed to be either functionally diverse or functionally simple based on belowground resource use patterns of constituent species. Communities were also constructed with different numbers of species (two or five) to examine interactions between species richness, functional diversity and invasion resistance. We hypothesized that communities with more complementary belowground resource use (i.e., more species rich and more functionally diverse communities) would be less easily invaded than communities with greater degrees of belowground resource use overlap. Two contrasting invasive species were introduced: an early-season, shallow rooting annual grass, Bromus hordeaceus (soft chess), and a late-season, deep rooting annual forb, Centaurea solstitialis (yellow starthistle). Invader responses to species richness and functional diversity treatments differed between sites. In general, the more similar the patterns of belowground resource use between residents of the plant community and the invader, the poorer the invader’s performance. Complementarity or overlap of resource use among species in the constructed communities appeared to affect invader success less than complementarity or overlap of resource use between the invader and the species present in the community.     

Functional and performance comparisons of invasive Hieracium lepidulum and co‐occurring species in New Zealand

Abstract One of the key environmental factors affecting plant species abundance, including that of invasive exotics, is nutrient resource availability. Plant functional response to nutrient availability, and what this tells us about plant interactions with associated species, may therefore give us clues about underlying processes related to plant abundance and invasion. Patterns of abundance of Hieracium lepidulum, a European herbaceous invader of subalpine New Zealand, appear to be related to soil fertility/nutrient availability, however, abundance may be influenced by other factors including disturbance. In this study we compare H. lepidulum and field co‐occurring species for growth performance across artificial nutrient concentration gradients, for relative competitiveness and for response to disturbance, to construct a functional profile of the species. Hieracium lepidulum was found to be significantly different in its functional response to nutrient concentration gradients. Hieracium lepidulum had high relative growth rate, high yield and root plasticity in response to nutrient concentration dilution, relatively low absolute yield, low competitive yield and a positive response to clipping disturbance relative to other species. Based on overall functional response to nutrient concentration gradients, compared with other species found at the same field sites, we hypothesize that H. lepidulum invasion is not related to competitive domination. Relatively low tolerance of nutrient dilution leads us to predict that H. lepidulum is likely to be restricted from invading low fertility sites, including sites within alpine vegetation or where intact high biomass plant communities are found. Positive response to clipping disturbance and relatively high nutrient requirement, despite poor competitive performance, leads us to predict that H. lepidulum may respond to selective grazing disturbance of associated vegetation. These results are discussed in relation to published observations of H. lepidulum in New Zealand and possible tests for the hypotheses raised here.     

Floral density and co-occurring congeners alter patterns of selection in annual plant communities*

Although the evolution and diversification of flowers is often attributed to pollinator-mediated selection, interactions between co-occurring plant species can alter patterns of selection mediated by pollinators and other agents. The extent to which both floral density and congeneric species richness affect patterns of net and pollinator-mediated selection on multiple co-occurring species in a community is unknown and is likely to depend on whether co-occurring plants experience competition or facilitation for reproduction. We conducted an observational study of selection on four species of Clarkia (Onagraceae) and tested for pollinator-mediated selection on two Clarkia species in communities differing in congeneric species richness and local floral density. When selection varied with community context, selection was generally stronger in communities with fewer species, where local conspecific floral density was higher, and where local heterospecific floral density was lower. These patterns suggest that intraspecific competition at high densities and interspecific competition at low densities may affect the evolution of floral traits. However, selection on floral traits was not pollinator mediated in Clarkia cylindrica or Clarkia xantiana, despite variation in pollinator visitation and the extent of pollen limitation across communities for C. cylindrica. As such, interactions between co-occurring species may alter patterns of selection mediated by abiotic agents of selection.     

Non-native grass invasion alters native plant composition in experimental communities

Invasions of non-native species are considered to have significant impacts on native species, but few studies have quantified the direct effects of invasions on native community structure and composition. Many studies on the effects of invasions fail to distinguish between (1) differential responses of native and non-native species to environmental conditions, and (2) direct impacts of invasions on native communities. In particular, invasions may alter community assembly following disturbance and prevent recolonization of native species. To determine if invasions directly impact native communities, we established 32 experimental plots (27.5 m²) and seeded them with 12 native species. Then, we added seed of a non-native invasive grass (Microstegium vimineum) to half of the plots and compared native plant community responses between control and invaded plots. Invasion reduced native biomass by 46, 64, and 58%, respectively, over three growing seasons. After the second year of the experiment, invaded plots had 43% lower species richness and 38% lower diversity as calculated from the Shannon index. Nonmetric multidimensional scaling ordination showed a significant divergence in composition between invaded and control plots. Further, there was a strong negative relationship between invader and native plant biomass, signifying that native plants are more strongly suppressed in densely invaded areas. Our results show that a non-native invasive plant inhibits native species establishment and growth following disturbance and that native species do not gain competitive dominance after multiple growing seasons. Thus, plant invaders can alter the structure of native plant communities and reduce the success of restoration efforts.     

Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra

Environmental gradients are caused by gradual changes in abiotic factors, which affect species abundances and distributions, and are important for the spatial distribution of biodiversity. One prominent environmental gradient is the altitude gradient. Understanding ecological processes associated with altitude gradients may help us to understand the possible effects climate change could have on species communities. We quantified vegetation cover, species richness, species evenness, beta diversity, and spatial patterns of community structure of vascular plants along altitude gradients in a subarctic mountain tundra in northern Sweden. Vascular plant cover and plant species richness showed unimodal relationships with altitude. However, species evenness did not change with altitude, suggesting that no individual species became dominant when species richness declined. Beta diversity also showed a unimodal relationship with altitude, but only for an intermediate spatial scale of 1 km. A lack of relationships with altitude for either patch or landscape scales suggests that any altitude effects on plant spatial heterogeneity occurred on scales larger than individual patches but were not effective across the whole landscape. We observed both nested and modular patterns of community structures, but only the modular patterns corresponded with altitude. Our observations point to biotic regulations of plant communities at high altitudes, but we found both scale dependencies and inconsistent magnitude of the effects of altitude on different diversity components. We urge for further studies evaluating how different factors influence plant communities in high altitude and high latitude environments, as well as studies identifying scale and context dependencies in any such influences.     

Species richness and evenness respond in a different manner to propagule density in developing prairie microcosm communities

Diversity has two basic components: richness, or number of species in a given area, and evenness, or how relative abundance or biomass is distributed among species. Previously, we found that richness and evenness can be negatively related across plant communities and that evenness can account for more variation in Shannon’s diversity index (H′) than richness, which suggests that relationships among diversity components can be complex. Non-positive relationships between evenness and richness could arise due to the effects of migration rate or local species interactions, and relationships could vary depending on how these two processes structure local communities. Here we test whether diversity components are equally or differentially affected over time by changes in seed density (and associated effects on established plant density and competition) in greenhouse communities during the very early stages of community establishment. In our greenhouse experiment, we seeded prairie microcosms filled with bare field soil at three densities with draws from a mix of 22 grass and forb species to test if increased competition intensity or seedling density would affect the relationships among diversity components during early community establishment. Increased seed density treatments caused diversity components to respond in a different manner and to have different relationships with time. Richness increased linearly with seed density early in the experiment when seedling emergence was high, but was unrelated to density later in the experiment. Evenness decreased log-linearly with seed densities on all sampling dates due to a greater dominance by Rudbeckia hirta with higher densities. Early in the experiment, diversity indices weakly reflected differences in richness, but later, after the competitive effects of Rudbeckia hirta became more intense, diversity indices more strongly reflected differences in evenness. This suggests that species evenness and diversity indices do not always positively covary with richness. Based on these results, we suggest that evenness and richness can be influenced by different processes, with richness being more influenced by the number of emerging seedlings and evenness more by species interactions like competition. These results suggest that both diversity components should be measured in plant diversity studies whenever it is possible.     

Biotic and abiotic constraints to a plant invasion in vegetation communities of Tierra del Fuego

The biotic resistance theory relates invader success to species richness, and predicts that, as species richness increases, invasibility decreases. The relationship between invader success and richness, however, seems to be positive at large scales of analysis, determined by abiotic constraints, and it is to be expected that it is negative at small scales, because of biotic interactions. Moreover, the negative relationship at small scales would be stronger within species of the same functional group, because of having similar resource exploitation mechanisms. We studied the relationship between the cover of a worldwide invader of grasslands, Hieracium pilosella L., and species richness, species diversity and the cover of different growth forms at two different levels of analysis in 128 sites during the initial invasion process in the Fuegian steppe, Southern Patagonia, Argentina. At regional level, the invader was positively correlated to total (r = 0.28, P = 0.003), exotic (r = 0.273, P = 0.004), and native species richness (r = 0.210, P = 0.026), and to species diversity (r = 0.193, P = 0.041). At community level, we found only a weak negative correlation between H. pilosella and total richness (r = ?0.426, P = 0.079) and diversity (r = ?0.658, P = 0.063). The relationship between the invader and other species of the same growth form was positive both at regional (r = 0.484, P < 0.001) and community (r = 0.593, P = 0.012) levels. Consequently, in the period of establishment and initial expansion of this exotic species, our results support the idea that invader success is related to abiotic factors at large scales of analysis. Also, we observed a possible sign of biotic constraint at community level, although this was not related to the abundance of species of the same growth form.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

An invasive plant provides refuge to native plant species in an intensely grazed ecosystem

Invasion by exotic plant species and herbivory can individually alter native plant species diversity, but their interactive effects in structuring native plant communities remain little studied. Many exotic plant species escape from their co-evolved specialized herbivores in their native range (in accordance with the enemy release hypothesis). When these invasive plants are relatively unpalatable, they may act as nurse plants by reducing herbivore damage on co-occurring native plants, thereby structuring native plant communities. However, the potential for unpalatable invasive plants to structure native plant communities has been little investigated. Here, we tested whether presence of an unpalatable exotic invader Opuntia ficus-indica was associated with the structure of native plant communities in an ecosystem with a long history of grazing by ungulate herbivores. Along 17 transects (each 1000 m long), we conducted a native vegetation survey in paired invaded and uninvaded plots. Plots that harboured O. ficus-indica had higher native plant species richness and Shannon–Wiener diversity H′ than uninvaded plots. However, mean species evenness J was similar between invaded and uninvaded plots. There was no significant correlation between native plant diversity and percentage plot cover by O. ficus-indica. Presence of O. ficus-indica was associated with a compositional change in native community assemblages between paired invaded and uninvaded plots. Although these results are only correlative, they suggest that unpalatable exotic plants may play an important ecological role as refugia for maintenance of native plant diversity in intensely grazed ecosystems.