Wildfire and cattle legacies on gradients of soil nitrogen underlie patterns of annual brome invasion.

Human activities are increasing wildfires and livestock activity in arid ecosystems with potential implications for the spread of invasive grasses. The objective of this study was to test whether fire history and cattle activity alter soil resource gradients, thereby affecting patterns of Bromus rubens L. (red brome) invasion. Six paired burned and unburned transect lines (1-km long) were established in the northeast Mojave Desert along the boundaries of four independent wildfire scars. At 100-m transect increment points, we measured the distance to the two nearest cowpats, and two random points and measured the density, height, biomass, and seed production of red brome, soil moisture and inorganic nitrogen (N). Cattle activity was 29% greater along burned transects compared to unburned transects (P < 0.05). Red brome height, density, and seed production were 11-34% greater along burned transects than unburned transects (P < 0.05). Red brome height, biomass, density, and seed production were twofold to tenfold greater next to cowpats compared to random points (P < 0.05). Soils along burned transects and beneath cowpats had greater soil inorganic N (P < 0.05), which was positively correlated with red brome density, height, biomass, and seed production (R2 = 0.60-0.85, P < 0.0001). Transgenerational effects were evident as seeds from red brome next to cowpats had 27% higher germination than seeds collected from random points. Positive responses of red brome to increased inorganic N related to fire and cattle activity may contribute fine fuel infill that drives invasive grass-fire cycles in deserts.

Ecosystem resilience to invasion and drought: Insights after 24 years in a rare never-grazed grassland.

Understanding the resilience of ecosystems globally is hampered by the complex and interacting drivers of change characteristic of the Anthropocene. This is true for drylands of the western US, where widespread alteration of disturbance regimes and spread of invasive non-native species occurred with westward expansion during the 1800s, including the introduction of domestic livestock and spread of Bromus tectorum, an invasive non-native annual grass. In addition, this region has experienced a multi-decadal drought not seen for at least 1200 years with potentially large and interacting impacts on native plant communities. Here, we present 24 years of twice-annual plant cover monitoring (1997-2021) from a semiarid grassland never grazed by domestic livestock but subject to a patchy invasion of B. tectorum beginning in ~1994, compare our findings to surveys done in 1967, and examine potential climate drivers of plant community changes. We found a significant warming trend in the study area, with more than 75% of study year temperatures being warmer than average (1966-2021). We observed a native perennial grass community with high resilience to climate forcings with cover values like those in 1967. In invaded patches, B. tectorum cover was greatest in the early years of this study (1997-2001; ~20%-40%) but was subsequently constrained by climate and subtle variation in soils, with limited evidence of long-term impacts to native vegetation, contradicting earlier studies. Our ability to predict year-to-year variation in functional group and species cover with climate metrics varied, with a 12-month integrated index and fall and winter patterns appearing most important. However, declines to near zero live cover in recent years in response to regional drought intensification leave questions regarding the resiliency of intact grasslands to ongoing aridification and whether the vegetation observations reported here may be a leading indicator of impending change in this protected ecosystem.

Marker-trait association analysis for drought tolerance in smooth bromegrass.

BACKGROUND: Little information is available on the application of marker-trait association (MTA) analysis for traits related to drought tolerance in smooth bromegrass. The objectives of this study were to identify marker loci associated with important agronomic traits and drought tolerance indices as well as fining stable associations in a diverse panel of polycross derived genotypes of smooth bromegrass. Phenotypic evaluations were performed at two irrigation regimes (normal and deficit irrigation) during 2 years; and association analysis was done with 626 SRAP markers. RESULTS: The results of population structure analysis identified three main subpopulations possessing significant genetic differences. Under normal irrigation, 68 and 57 marker-trait associations were identified using general linear model (GLM) and mixed linear mode1 (MLM), respectively. While under deficit irrigation, 61 and 54 markers were associated with the genes controlling the studied traits, based on these two models, respectively. Some of the markers were associated with more than one trait. It was revealed that markers Me1/Em5-11, Me1/Em3-15, and Me5/Em4-7 were consistently linked with drought-tolerance indices. CONCLUSION: Following marker validation, the MTAs reported in this panel could be useful tools to initiate marker-assisted selection (MAS) and targeted trait introgression of smooth bromegrass under normal and deficit irrigation regimes, and possibly fine mapping and cloning of the underlying genes and QTLs.

Livestock grazing is associated with seasonal reduction in pollinator biodiversity and functional dispersion but cheatgrass invasion is not: Variation in bee assemblages in a multi-use shortgrass prairie.

Livestock grazing and non-native plant species affect rangeland habitats globally. These factors may have important effects on ecosystem services including pollination, yet, interactions between pollinators, grazing, and invasive plants are poorly understood. To address this, we tested the hypothesis that cattle grazing and site colonization by cheatgrass (Bromus tectorum) impact bee foraging and nesting habitats, and the biodiversity of wild bee communities, in a shortgrass prairie system. Bee nesting habitats (litter and wood cover) were marginally improved in non-grazed sites with low cheatgrass cover, though foraging habitat (floral cover and richness, bare soil) did not differ among cattle-grazed sites or non-grazed sites with low or high cheatgrass cover. However, floral cover was a good predictor of bee abundance and functional dispersion. Mean bee abundance, richness, diversity and functional diversity were significantly lower in cattle-grazed habitats than in non-grazed habitats. Differences in bee diversity among habitats were pronounced early in the growing season (May) but by late-season (August) these differences eroded as Melissodes spp. and Bombus spp. became more abundant at study sites. Fourth-corner analysis revealed that sites with high floral cover tended to support large, social, polylectic bees; sites with high grass cover tended to support oligolectic solitary bees. Both cattle-grazed sites and sites with high cheatgrass cover were associated with lower abundances of above-ground nesting bees but higher abundance of below-ground nesters than non-grazed sites with low cheatgrass cover. We conclude that high cheatgrass cover is not associated with reduced bee biodiversity or abundance, but cattle grazing was negatively associated with bee abundances and altered species composition. Although floral cover is an important predictor of bee assemblages, this was not impacted by cattle grazing and our study suggests that cattle likely impact bee communities through effects other than those mediated by forbs, including soil disturbance or nest destruction. Efforts aimed at pollinator conservation in prairie habitats should focus on managing cattle impacts early in the growing season to benefit sensitive bee species.

The effect of consumer pressure and abiotic stress on positive plant interactions are mediated by extreme climatic events.

Environmental extremes resulting from a changing climate can have profound implications for plant interactions in desert communities. Positive interactions can buffer plant communities from abiotic stress and consumer pressure caused by climatic extremes, but limited research has explored this empirically. We tested the hypothesis that the mechanism of shrub facilitation on an annual plant community can change with precipitation extremes in deserts. During years of extreme drought and above-average rainfall in a desert, we measured plant interactions and biomass while manipulating a soil moisture gradient and reducing consumer pressure. Shrubs facilitated the annual plant community at all levels of soil moisture through reductions in microclimatic stress in both years and herbivore protection in the wet year only. Shrub facilitation and the high rainfall year contributed to the dominance of a competitive annual species in the plant community. Precipitation patterns in deserts determine the magnitude and type of facilitation mechanisms. Moreover, shrub facilitation mediates the interspecific competition within the associated annual community between years with different rainfall amounts. Examining multiple drivers during extreme climate events is a challenging area of research, but it is a necessary consideration given forecasts predicting that these events will increase in frequency and magnitude.

Novel fine-scale aerial mapping approach quantifies grassland weed cover dynamics and response to management.

Invasive weeds threaten the biodiversity and forage productivity of grasslands worldwide. However, management of these weeds is constrained by the practical difficulty of detecting small-scale infestations across large landscapes and by limits in understanding of landscape-scale invasion dynamics, including mechanisms that enable patches to expand, contract, or remain stable. While high-end hyperspectral remote sensing systems can effectively map vegetation cover, these systems are currently too costly and limited in availability for most land managers. We demonstrate application of a more accessible and cost-effective remote sensing approach, based on simple aerial imagery, for quantifying weed cover dynamics over time. In California annual grasslands, the target communities of interest include invasive weedy grasses (Aegilops triuncialis and Elymus caput-medusae) and desirable forage grass species (primarily Avena spp. and Bromus spp.). Detecting invasion of annual grasses into an annual-dominated community is particularly challenging, but we were able to consistently characterize these two communities based on their phenological differences in peak growth and senescence using maximum likelihood supervised classification of imagery acquired twice per year (in mid- and end-of season). This approach permitted us to map weed-dominated cover at a 1-m scale (correctly detecting 93% of weed patches across the landscape) and to evaluate weed cover change over time. We found that weed cover was more pervasive and persistent in management units that had no significant grazing for several years than in those that were grazed, whereas forage cover was more abundant and stable in the grazed units. This application demonstrates the power of this method for assessing fine-scale vegetation transitions across heterogeneous landscapes. It thus provides means for small-scale early detection of invasive species and for testing fundamental questions about landscape dynamics.

A warmer and drier climate in the northern sagebrush biome does not promote cheatgrass invasion or change its response to fire.

Dryland shrub communities have been degraded by a range of disturbances and now face additional stress from global climate change. The spring/summer growing season of the North American sagebrush biome is projected to become warmer and drier, which is expected to facilitate the expansion of the invasive annual grass Bromus tectorum (cheatgrass) and alter its response to fire in the northern extent of the biome. We tested these predictions with a factorial experiment with two levels of burning (spring burn and none) and three climate treatments (warming, warming + drying, and control) that was repeated over 3 years in a Montana sagebrush steppe. We expected the climate treatments to make B. tectorum more competitive with the native perennial grass community, especially Pseudoroegneria spicata, and alter its response to fire. Experimental warming and warming + drying reduced B. tectorum cover, biomass, and fecundity, but there was no response to fire except for fecundity, which increased; the native grass community was the most significant factor that affected B. tectorum metrics. The experimental climate treatments also negatively affected P. spicata, total native grass cover, and community biodiversity, while fire negatively affected total native grass cover, particularly when climate conditions were warmer and drier. Our short-term results indicate that without sufficient antecedent moisture and a significant disruption to the native perennial grass community, a change in climate to a warmer and drier spring/summer growing season in the northern sagebrush biome will not facilitate B. tectorum invasion or alter its response to fire.

Species' traits help predict small mammal responses to habitat homogenization by an invasive grass.

Invasive plants can negatively affect native species, however, the strength, direction, and shape of responses may vary depending on the type of habitat alteration and the natural history of native species. To prioritize conservation of vulnerable species, it is therefore critical to effectively predict species' responses to invasive plants, which may be facilitated by a framework based on species' traits. We studied the population and community responses of small mammals and changes in habitat heterogeneity across a gradient of cheatgrass (Bromus tectorum) cover, a widespread invasive plant in North America. We live-trapped small mammals over two summers and assessed the effect of cheatgrass on native small mammal abundance, richness, and species-specific and trait-based occupancy, while accounting for detection probability and other key habitat elements. Abundance was only estimated for the most common species, deer mice (Peromyscus maniculatus). All species were pooled for the trait-based occupancy analysis to quantify the ability of small mammal traits (habitat association, mode of locomotion, and diet) to predict responses to cheatgrass invasion. Habitat heterogeneity decreased with cheatgrass cover. Deer mouse abundance increased marginally with cheatgrass. Species richness did not vary with cheatgrass, however, pocket mouse (Perognathus spp.) and harvest mouse (Reithrodontomys spp.) occupancy tended to decrease and increase, respectively, with cheatgrass cover, suggesting a shift in community composition. Cheatgrass had little effect on occupancy for deer mice, 13-lined ground squirrels (Spermophilus tridecemlineatus), and Ord's kangaroo rat (Dipodomys ordii). Species' responses to cheatgrass primarily corresponded with our a priori predictions based on species' traits. The probability of occupancy varied significantly with a species' habitat association but not with diet or mode of locomotion. When considered within the context of a rapid habitat change, such as caused by invasive plants, relevant species' traits may provide a useful framework for predicting species' responses to a variety of habitat disturbances. Understanding which species are likely to be most affected by exotic plant invasion will help facilitate more efficient, targeted management and conservation of native species and habitats.

Native fungal endophytes suppress an exotic dominant and increase plant diversity over small and large spatial scales.

Understanding community dynamics and processes, such as the factors that generate and maintain biodiversity, drive succession, and affect invasion susceptibility, is a central goal in ecology and evolution. While most studies of how species interactions affect communities have focused on highly visible macroorganisms, we show that mutualistic microfungal endophytes have community-level effects across their host plant's range and provide the first example of fungal endophytes enhancing plant diversity. A three-year field study in which we experimentally manipulated endophyte abundance in a native Californian grass showed that despite their minute biomass, endophytes dramatically increased plant community diversity (~110% greater increase with endophytes) by suppressing a dominant invasive grass, Bromus diandrus. This effect was also detectable, but smaller, across five additional common gardens spanning ecologically diverse habitats, different climates, and > 400 km of the host grass' range as well as at microspatial scales within gardens. Our study illustrates that mutualistic microbes, while often hidden players, can have unexpectedly large ecological impacts across a wide range of habitats and scales and may be important for promoting diverse communities and ecosystems.

EPSPS gene amplification in glyphosate-resistant Bromus diandrus.

BACKGROUND: Glyphosate is the most widely used herbicide in the world and has been intensively used to control B. diandrus, a problematic weed of crops and pastures in southern Australia. RESULTS: Resistance to glyphosate was identified in two populations of B. diandrus that were nearly fivefold more resistant to glyphosate than wild-type plants. Both populations contained EPSPS gene amplification, with resistant plants having an average of around 20-fold the number of copies of EPSPS compared with susceptible plants. EPSPS expression was also increased in resistant plants of both populations; however, expression levels were not correlated with the number of EPSPS copies. Amplification of only one of the four EPSPS genes present in B. diandus was detected. Investigation into the inheritance of glyphosate resistance found no segregation in the F2 generation. Every individual in the F2 populations contained between three and 30 copies of EPSPS; however, on average they contained fewer copies compared with the parent resistant population. CONCLUSIONS: Glyphosate resistance in B. diandrus is due to EPSPS gene amplification. Resistance is heritable but complex.

Effects of precipitation change and neighboring plants on population dynamics of Bromus tectorum.

Shifting precipitation patterns resulting from global climate change will influence the success of invasive plant species. In the Front Range of Colorado, Bromus tectorum (cheatgrass) and other non-native winter annuals have invaded grassland communities and are becoming more abundant. As the global climate warms, more precipitation may fall as rain rather than snow in winter, and an increase in winter rain could benefit early-growing winter annuals, such as B. tectorum, to the detriment of native species. In this study we measured the effects of simulated changes in seasonal precipitation and presence of other plant species on population growth of B. tectorum in a grassland ecosystem near Boulder, Colorado, USA. We also performed elasticity analyses to identify life transitions that were most sensitive to precipitation differences. In both study years, population growth rates were highest for B. tectorum growing in treatments receiving supplemental winter precipitation and lowest for those receiving the summer drought treatment. Survival of seedlings to flowering and seed production contributed most to population growth in all treatments. Biomass of neighboring native plants was positively correlated with reduced population growth rates of B. tectorum. However, exotic plant biomass had no effect on population growth rates. This study demonstrates how interacting effects of climate change and presence of native plants can influence the population growth of an invasive species. Overall, our results suggest that B. tectorum will become more invasive in grasslands if the seasonality of precipitation shifts towards wetter winters and allows B. tectorum to grow when competition from native species is low.

Interacting agricultural pests and their effect on crop yield: application of a Bayesian decision theory approach to the joint management of Bromus tectorum and Cephus cinctus.

Worldwide, the landscape homogeneity of extensive monocultures that characterizes conventional agriculture has resulted in the development of specialized and interacting multitrophic pest complexes. While integrated pest management emphasizes the need to consider the ecological context where multiple species coexist, management recommendations are often based on single-species tactics. This approach may not provide satisfactory solutions when confronted with the complex interactions occurring between organisms at the same or different trophic levels. Replacement of the single-species management model with more sophisticated, multi-species programs requires an understanding of the direct and indirect interactions occurring between the crop and all categories of pests. We evaluated a modeling framework to make multi-pest management decisions taking into account direct and indirect interactions among species belonging to different trophic levels. We adopted a Bayesian decision theory approach in combination with path analysis to evaluate interactions between Bromus tectorum (downy brome, cheatgrass) and Cephus cinctus (wheat stem sawfly) in wheat (Triticum aestivum) systems. We assessed their joint responses to weed management tactics, seeding rates, and cultivar tolerance to insect stem boring or competition. Our results indicated that C. cinctus oviposition behavior varied as a function of B. tectorum pressure. Crop responses were more readily explained by the joint effects of management tactics on both categories of pests and their interactions than just by the direct impact of any particular management scheme on yield. In accordance, a C. cinctus tolerant variety should be planted at a low seeding rate under high insect pressure. However as B. tectorum levels increase, the C. cinctus tolerant variety should be replaced by a competitive and drought tolerant cultivar at high seeding rates despite C. cinctus infestation. This study exemplifies the necessity of accounting for direct and indirect biological interactions occurring within agroecosystems and propagating this information from the statistical analysis stage to the management stage.

Using high-resolution future climate scenarios to forecast Bromus tectorum invasion in Rocky Mountain National Park.

National Parks are hallmarks of ecosystem preservation in the United States. The introduction of alien invasive plant species threatens protection of these areas. Bromus tectorum L. (commonly called downy brome or cheatgrass), which is found in Rocky Mountain National Park (hereafter, the Park), Colorado, USA, has been implicated in early spring competition with native grasses, decreased soil nitrogen, altered nutrient and hydrologic regimes, and increased fire intensity. We estimated the potential distribution of B. tectorum in the Park based on occurrence records (n = 211), current and future climate, and distance to roads and trails. An ensemble of six future climate scenarios indicated the habitable area of B. tectorum may increase from approximately 5.5% currently to 20.4% of the Park by the year 2050. Using ordination methods we evaluated the climatic space occupied by B. tectorum in the Park and how this space may shift given future climate change. Modeling climate change at a small extent (1,076 km2) and at a fine spatial resolution (90 m) is a novel approach in species distribution modeling, and may provide inference for microclimates not captured in coarse-scale models. Maps from our models serve as high-resolution hypotheses that can be improved over time by land managers to set priorities for surveys and removal of invasive species such as B. tectorum.

Plastic responses of native plant root systems to the presence of an invasive annual grass.

UNLABELLED: • PREMISE OF THE STUDY: The ability to respond to environmental change via phenotypic plasticity may be important for plants experiencing disturbances such as climate change and plant invasion. Responding to belowground competition through root plasticity may allow native plants to persist in highly invaded systems such as the cold deserts of the Intermountain West, USA.• METHODS: We investigated whether Poa secunda, a native bunchgrass, could alter root morphology in response to nutrient availability and the presence of a competitive annual grass. Seeds from 20 families were grown with high and low nutrients and harvested after 50 d, and seeds from 48 families, grown with and without Bromus tectorum, were harvested after ∼2 or 6 mo. We measured total biomass, root mass fraction, specific root length (SRL), root tips, allocation to roots of varying diameter, and plasticity in allocation.• KEY RESULTS: Plants had many parallel responses to low nutrients and competition, including increased root tip production, a trait associated with tolerance to reduced resources, though families differed in almost every trait and correlations among trait changes varied among experiments, indicating flexibility in plant responses. Seedlings actively increased SRL and fine root allocation under competition, while older seedlings also increased coarse root allocation, a trait associated with increased tolerance, and increased root mass fraction.• CONCLUSIONS: The high degree of genetic variation for root plasticity within natural populations could aid in the long-term persistence of P. secunda because phenotypic plasticity may allow native species to persist in invaded and fluctuating resource environments.

Contemporary evolution of an invasive grass in response to elevated atmospheric CO(2) at a Mojave Desert FACE site.

Elevated atmospheric CO2 has been shown to rapidly alter plant physiology and ecosystem productivity, but contemporary evolutionary responses to increased CO2 have yet to be demonstrated in the field. At a Mojave Desert FACE (free-air CO2 enrichment) facility, we tested whether an annual grass weed (Bromus madritensis ssp. rubens) has evolved in response to elevated atmospheric CO2 . Within 7 years, field populations exposed to elevated CO2 evolved lower rates of leaf stomatal conductance; a physiological adaptation known to conserve water in other desert or water-limited ecosystems. Evolution of lower conductance was accompanied by reduced plasticity in upregulating conductance when CO2 was more limiting; this reduction in conductance plasticity suggests that genetic assimilation may be ongoing. Reproductive fitness costs associated with this reduction in phenotypic plasticity were demonstrated under ambient levels of CO2 . Our findings suggest that contemporary evolution may facilitate this invasive species' spread in this desert ecosystem.

Long-term response of a Mojave Desert winter annual plant community to a whole-ecosystem atmospheric CO2 manipulation (FACE).

Desert annuals are a critically important component of desert communities and may be particularly responsive to increasing atmospheric (CO2 ) because of their high potential growth rates and flexible phenology. During the 10-year life of the Nevada Desert FACE (free-air CO2 enrichment) Facility, we evaluated the productivity, reproductive allocation, and community structure of annuals in response to long-term elevated (CO2 ) exposure. The dominant forb and grass species exhibited accelerated phenology, increased size, and higher reproduction at elevated (CO2 ) in a wet El Niño year near the beginning of the experiment. However, a multiyear dry cycle resulted in no increases in productivity or reproductive allocation for the remainder of the experiment. At the community level, early indications of increased dominance of the invasive Bromus rubens at elevated (CO2 ) gave way to an absence of Bromus in the community during a drought cycle, with a resurgence late in the experiment in response to higher rainfall and a corresponding high density of Bromus in a final soil seed bank analysis, particularly at elevated (CO2 ). This long-term experiment resulted in two primary conclusions: (i) elevated (CO2 ) does not increase productivity of annuals in most years; and (ii) relative stimulation of invasive grasses will likely depend on future precipitation, with a wetter climate favoring invasive grasses but currently predicted greater aridity favoring native dicots.

Abscisic acid induced freezing tolerance in chilling-sensitive suspension cultures and seedlings of rice.

BACKGROUND: The role of abscisic acid (ABA) as a possible activator of cold acclimation process was postulated since endogenous levels of ABA increase temporarily or constitutively during cold-hardening. Exogenous application of ABA has been known to induce freezing tolerance at ambient temperatures in in vitro systems derived from cold hardy plants. Yet, some cell cultures acquired much greater freezing tolerance by ABA than by cold whilst maintaining active growth. This raises questions about the relationships among ABA, cold acclimation and growth cessation. To address this question, we attempted to 1) determine whether exogenous ABA can confer freezing tolerance in chilling-sensitive rice suspension cells and seedlings, which obviously lack the mechanisms to acquire freezing tolerance in response to cold; 2) characterize this phenomenon by optimizing the conditions and compare with the case of cold hardy bromegrass cells. RESULTS: Non-embryogenic suspension cells of rice suffered serious chilling injury when exposed to 4°C. When incubated with ABA at the optimal conditions (0.5-1 g cell inoculum, 75 µM ABA, 25-30°C, 7-10 days), they survived slow freezing (2°C/h) to -9.0 ~ -9.3°C (LT50: 50% killing temperature) while control cells were mostly injured at -3°C (LT50: -0.5 ~ -1.5°C). Ice-inoculation of the cell suspension at -3°C and survival determination by regrowth confirmed that ABA-treated rice cells survived extracellular freezing at -9°C. ABA-induced freezing tolerance did not require any exposure to cold and was best achieved at 25-30°C where the rice cells maintained high growth even in the presence of ABA. ABA treatment also increased tolerance to heat (43°C) as determined by regrowth. ABA-treated cells tended to have more augmented cytoplasm and/or reduced vacuole sizes compared to control cultures with a concomitant increase in osmolarity and a decrease in water content. ABA-treated (2-7 days) in vitro grown seedlings and their leaves survived slow freezing to -3°C with only marginal injury (LT50: -4°C) whereas untreated seedlings were killed at -3°C (LT50: -2°C). CONCLUSIONS: The results indicate that exogenous ABA can induce some levels of freezing tolerance in chilling-sensitive rice cells and seedlings, probably by eliciting mechanisms different from low temperature-induced cold acclimation.

A road map for integrating eco-evolutionary processes into biodiversity models.

The demand for projections of the future distribution of biodiversity has triggered an upsurge in modelling at the crossroads between ecology and evolution. Despite the enthusiasm around these so-called biodiversity models, most approaches are still criticised for not integrating key processes known to shape species ranges and community structure. Developing an integrative modelling framework for biodiversity distribution promises to improve the reliability of predictions and to give a better understanding of the eco-evolutionary dynamics of species and communities under changing environments. In this article, we briefly review some eco-evolutionary processes and interplays among them, which are essential to provide reliable projections of species distributions and community structure. We identify gaps in theory, quantitative knowledge and data availability hampering the development of an integrated modelling framework. We argue that model development relying on a strong theoretical foundation is essential to inspire new models, manage complexity and maintain tractability. We support our argument with an example of a novel integrated model for species distribution modelling, derived from metapopulation theory, which accounts for abiotic constraints, dispersal, biotic interactions and evolution under changing environmental conditions. We hope such a perspective will motivate exciting and novel research, and challenge others to improve on our proposed approach.

Global change effects on Bromus tectorum L. (Poaceae) at its high-elevation range margin.

Global change is likely to affect invasive species distribution, especially at range margins. In the eastern Sierra Nevada, California, USA, the invasive annual grass, Bromus tectorum, is patchily distributed and its impacts have been minimal compared with other areas of the Intermountain West. We used a series of in situ field manipulations to determine how B. tectorum might respond to changing climatic conditions and increased nitrogen deposition at the high-elevation edge of its invaded range. Over 3 years, we used snow fences to simulate changes in snowpack, irrigation to simulate increased frequency and magnitude of springtime precipitation, and added nitrogen (N) at three levels (0, 5, and 10 g m(-2) ) to natural patches of B. tectorum growing under the two dominant shrubs, Artemisia tridentata and Purshia tridentata, and in intershrub spaces (INTR). We found that B. tectorum seedling density in April was lower following deeper snowpack possibly due to delayed emergence, yet there was no change in spikelet production or biomass accumulation at the time of harvest. Additional spring rain events increased B. tectorum biomass and spikelet production in INTR plots only. Plants were primarily limited by water in 2009, but colimited by N and water in 2011, possibly due to differences in antecedent moisture conditions at the time of treatments. The threshold at which N had an effect varied with magnitude of water additions. Frequency of rain events was more influential than magnitude in driving B. tectorum growth and fecundity responses. Our results suggest that predicted shifts from snow to rain could facilitate expansion of B. tectorum at high elevation depending on timing of rain events and level of N deposition. We found evidence for P-limitation at this site and an increase in P-availability with N additions, suggesting that stoichiometric relationships may also influence B. tectorum spread.

Land use causes genetic differentiation of life-history traits in Bromus hordeaceus.

There is increasing evidence that species can evolve rapidly in response to environmental change. However, although land use is one of the key drivers of current environmental change, studies of its evolutionary consequences are still fairly scarce, in particular studies that examine land-use effects across large numbers of populations, and discriminate between different aspects of land use. Here, we investigated genetic differentiation in relation to land use in the annual grass Bromus hordeaceus. A common garden study with offspring from 51 populations from three regions and a broad range of land-use types and intensities showed that there was indeed systematic population differentiation of ecologically important plant traits in relation to land use, in particular due to increasing mowing and grazing intensities. We also found strong land-use-related genetic differentiation in plant phenology, where the onset of flowering consistently shifted away from the typical time of management. In addition, increased grazing intensity significantly increased the genetic variability within populations. Our study suggests that land use can cause considerable genetic differentiation among plant populations, and that the timing of land use may select for phenological escape strategies, particularly in monocarpic plant species.