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Plant community resistance to invasion by Bromus species: The roles of community attributes, Bromus interactions with plant communities, and Bromus traits

The factors that determine plant community resistance to exotic annual Bromus species ( Bromus hereafter) are diverse and context specific. They are influenced by the environmental characteristics and attributes of the community, the traits of Bromus species, and the direct and indirect interactions of Bromus with the plant community. Environmental factors, in particular ambient and soil temperatures, have significant effects on the ability of Bromus to establish and spread. Seasonality of precipitation relative to temperature influences plant community resistance to Bromus through effects on soil water storage, timing of water and nutrient availability, and dominant plant life forms. Differences among plant communities in how well soil resource use by the plant community matches resource supply rates can influence the magnitude of resource fluctuations due to either climate or disturbance and thus the opportunities for invasion. The spatial and temporal patterns of resource availability and acquisition of growth resources by Bromus versus native species strongly influence resistance to invasion. Traits of Bromus that confer a “priority advantage” for resource use in many communities include early-season germination and high growth and reproductive rates. Resistance to Bromus can be overwhelmed by high propagule supply, low innate seed dormancy, and large, if short-lived, seed banks. Biological crusts can inhibit germination and establishment of invasive annual plants, including several annual Bromus species, but are effective only in the absence of disturbance. Herbivores can have negative direct effects on Bromus , but positive indirect effects through decreases in competitors. Management strategies can be improved through increased understanding of community resistance to exotic annual Bromus species.

Book chapter↗

Land uses, fire, and invasion: Exotic annual Bromus and human dimensions

Human land uses are the primary cause of the introduction and spread of exotic annual Bromus species. Initial introductions were likely linked to contaminated seeds used by homesteading farmers in the late 1880s and early 1900s. Transportation routes aided their spread. Unrestricted livestock grazing from the 1800s through the mid-1900s reduced native plant competitors leaving large areas vulnerable to Bromus dominance. Ecosystems with cooler and moister soils tend to have greater potential to recover from disturbances (resilience) and to be more resistant to Bromus invasion and dominance. Warmer and drier ecosystems are less resistant to Bromus and are threatened by altered fire regimes which can lead to Bromus dominance, impacts to wildlife, and alternative stable states. Native Americans used fire for manipulating plant communities and may have contributed to the early dominance of Bromus in portions of California. Fire as a tool is now limited to site preparation for revegetation in most ecosystems where Bromus is a significant problem. Once Bromus dominates, breaking annual grass/fire cycles requires restoring fire-tolerant perennial grasses and forbs, which can compete with Bromus and resist its dominance. Current weed management policies often lack regulations to prevent further expansion of Bromus . Research is needed on how and where livestock grazing might help increase perennial grass and forb cover and density to create ecosystems that are more resistant to Bromus . Also, studies are needed to ascertain the role, if any, of oil and gas development in contributing to the spread of Bromus .

Book chapter↗

Exotic annual Bromus invasions: Comparisons among species and ecoregions in the western United States

Exotic annual Bromus species are widely recognized for their potential to invade, dominate, and alter the structure and function of ecosystems. In this chapter, we summarize the invasion potential, ecosystem threats, and management strategies for different Bromus species within each of five ecoregions of the western United States. We characterize invasion potential and threats in terms of ecosystem resistance to Bromus invasion and ecosystem resilience to disturbance with an emphasis on the importance of fire regimes. We also explain how soil temperature and moisture regimes can be linked to patterns of resistance and resilience and provide a conceptual framework that can be used to evaluate the relative potential for invasion and ecological impact of the dominant exotic annual Bromus species in the western United States.

Book chapter↗

Ecosystem impacts of exotic annual invaders in the genus Bromus

An understanding of the impacts of exotic plant species on ecosystems is necessary to justify and guide efforts to limit their spread, restore natives, and plan for conservation. Invasive annual grasses such as Bromus tectorum , B. rubens , B. hordeaceus , and B. diandrus (hereafter collectively referred to as Bromus ) transform the structure and function of ecosystems they dominate. Experiments that prove cause-and-effect impacts of Bromus are rare, yet inferences can be gleaned from the combination of Bromus -ecosystem associations, ecosystem condition before/after invasion, and an understanding of underlying mechanisms. Bromus typically establishes in bare soil patches and can eventually replace perennials such as woody species or bunchgrasses, creating a homogeneous annual cover. Plant productivity and cover are less stable across seasons and years when Bromus dominates, due to a greater response to annual climate variability. Bromus’ “flash” of growth followed by senescence early in the growing season, combined with shallow rooting and annual habit, may lead to incomplete use of deep soil water, reduced C sequestration, and accelerated nutrient cycling. Litter produced by Bromus alters nearly all aspects of ecosystems and notably increases wildfire occurrence. Where Bromus has become dominant, it can decrease soil stability by rendering soils bare for months following fire or episodic, pathogen-induced stand failure. Bromus-invaded communities have lower species diversity, and associated species tend to be generalists adapted to unstable and variable habitats. Changes in litter, fire, and soil properties appear to feedback to reinforce Bromus’ dominance in a pattern that portends desertification.

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Soil moisture and biogeochemical factors influence the distribution of annual Bromus species

Abiotic factors have a strong influence on where annual Bromus species are found. At the large regional scale, temperature and precipitation extremes determine the boundaries of Bromus occurrence. At the more local scale, soil characteristics and climate influence distribution, cover, and performance. In hot, dry, summer-rainfall-dominated deserts (Sonoran, Chihuahuan), little or no Bromus is found, likely due to timing or amount of soil moisture relative to Bromus phenology. In hot, winter-rainfall-dominated deserts (parts of the Mojave Desert), Bromus rubens is widespread and correlated with high phosphorus availability. It also responds positively to additions of nitrogen alone or with phosphorus. On the Colorado Plateau, with higher soil moisture availability, factors limiting Bromus tectorum populations vary with life stage: phosphorus and water limit germination, potassium and the potassium/magnesium ratio affect winter performance, and water and potassium/magnesium affect spring performance. Controlling nutrients also change with elevation. In cooler deserts with winter precipitation (Great Basin, Columbia Plateau) and thus even greater soil moisture availability, B. tectorum populations are controlled by nitrogen, phosphorus, or potassium. Experimental nitrogen additions stimulate Bromus performance. The reason for different nutrients limiting in dissimilar climatic regions is not known, but it is likely that site conditions such as soil texture (as it affects water and nutrient availability), organic matter, and/or chemistry interact in a manner that regulates nutrient availability and limitations. Under future drier, hotter conditions, Bromus distribution is likely to change due to changes in the interaction between moisture and nutrient availability.

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Invertebrates in managed waterfowl marshes

Invertebrates are an important food for breeding, migrating, and wintering waterfowl. Sparse study has been devoted to understanding the influence of waterfowl and wetland management on production of invertebrates for waterfowl foods; however, manipulation of hydrology and soils may change or enhance production. Fish can compete with waterfowl for invertebrate forage in wetlands and harm aquatic macrophytes; biomanipulation (e.g., stocking piscivores) may improve waterfowl habitat quality. Similarly, some terrestrial vertebrates (e.g., beaver ( Castor canadensis )) may positively or negatively impact invertebrate communities in waterfowl habitats. Various challenges exist to wetland management for invertebrates for waterfowl, but the lack of data on factors influencing production may be the most limiting.

Book chapter↗

Participatory modeling and structured decision making

Structured decision making (SDM) provides a framework for making sound decisions even when faced with uncertainty, and is a transparent, defensible, and replicable method used to understand complex problems. A hallmark of SDM is the explicit incorporation of values and science, which often includes participation from multiple stakeholders, helping to garner trust and ultimately result in a decision that is more likely to be implemented. The core steps in the SDM process are used to structure thinking about natural resources management choices, and include: (1) properly defining the problem and the decision context, (2) determining the objectives that help describe the aspirations of the decision maker, (3) devising management actions or alternatives that can achieve those objectives, (4) evaluating the outcomes or consequences of each alternative on each of the objectives, (5) evaluating trade-offs, and (6) implementing the decision. Participatory modeling for SDM includes engaging stakeholders in some or all of the steps of the SDM process listed above. In addition, participatory modeling often is crucial for creating qualitative and quantitative models of how the system works, providing data for these models, and eliciting expert opinion when data are unavailable. In these ways, SDM provides a framework for decision making in natural resources management that includes participation from stakeholder groups throughout the process, including the modeling phase.

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Case studies of capacity building for biodiversity monitoring

Monitoring the status and trends of species is critical to their conservation and management. However, the current state of biodiversity monitoring is insufficient to detect such for most species and habitats, other than in a few localised areas. One of the biggest obstacles to adequate monitoring is the lack of local capacity to carry out such programs. Thus, building the capacity to do such monitoring is imperative. We here highlight different biodiversity monitoring efforts to illustrate how capacity building efforts are being conducted at different geographic scales and under a range of resource, literacy, and training constraints. Accordingly, we include examples of monitoring efforts from within countries (Kenya, France, and China), within regions (Central America and the Arctic) and larger capacity building programs including EDGE (Evolutionarily Distinct and Globally Endangered) of Existence and the National Red List Alliance.

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The physiology of mangrove trees with changing climate

Mangrove forests grow on saline, periodically flooded soils of the tropical and subtropical coasts. The tree species that comprise the mangrove are halophytes that have suites of traits that confer differing levels of tolerance of salinity, aridity, inundation and extremes of temperature. Here we review how climate change and elevated levels of atmospheric CO2 will influence mangrove forests. Tolerance of salinity and inundation in mangroves is associated with the efficient use of water for photosynthetic carbon gain which unpins anticipated gains in productivity with increasing levels of CO2. We review evidence of increases in productivity with increasing CO2, finding that enhancements in growth appear to be similar to trees in non-mangrove habitats and that gains in productivity with elevated CO2 are likely due to changes in biomass allocation. High levels of trait plasticity are observed in some mangrove species, which potentially facilitates their responses to climate change. Trait plasticity is associated with broad tolerance of salinity, aridity, low temperatures and nutrient availability. Because low temperatures and aridity place strong limits on mangrove growth at the edge of their current distribution, increasing temperatures over time and changing rainfall patterns are likely to have an important influence on the distribution of mangroves. We provide a global analysis based on plant traits and IPCC scenarios of changing temperature and aridity that indicates substantial global potential for mangrove expansion.

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Biological soil crusts as an organizing principle in drylands

Biological soil crusts (biocrusts) have been present on Earth’s terrestrial surfaces for billions of years. They are a critical part of ecosystem processes in dryland regions, as they cover most of the soil surface and thus mediate almost all inputs and outputs from soils in these areas. There are many intriguing, but understudied, roles these communities may play in drylands. These include their function in nutrient capture and transformation, influence on the movement and distribution of nutrients and water within dryland soils, ability to structure vascular plant communities, role in creating biodiversity hotspots, and the possibility that they can be used as indicators of soil health. There are still many fascinating aspects of these communities that need study, and we hope that this chapter will facilitate such efforts.

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Controls on distribution patterns of biological soil crusts at micro- to global scales

Biocrusts are heterogeneously distributed in space. The drivers of their distribution patterns vary depending on the spatial scale of observation. Globally, there are about 1337 cyanobacteria, algae, bryophyte, and lichen species reported as components of biocrusts. At the broadest biogeographical scales, the degree and age of isolation of land masses may dictate distribution of these species and the similarities of the floras of different continents. At intra-continental and smaller scales, climate strongly influences abundance and community composition of biocrusts. Within drylands, biocrusts become more abundant as precipitation increases. The seasonality of rainfall is about equally important, with regions receiving most precipitation as winter rain and snow exhibiting the highest abundance and greatest relative cover of bryophyte and lichens vs. cyanobacteria. Temperature gradients may dictate the dominant cyanobacterium present in the community. At eco-regional and smaller scales, edaphic gradients determined by either soil parent materials or geomorphology or both become particularly influential. Globally, the most significant soil properties influencing the eco-regional scale cover and richness of biocrusts in dryland environments are soil texture, pH, and soil CaCO3 content. Sandier soils tend to favour development of cyanobacterial biocrusts, whereas mosses and lichens tend to be more abundant on finer textured soils. The alkalinity and CaCO3 content of soils are associated with greater bioocrust abundance in some regions, and dictates the species composition in the bryophyte and lichen component. Globally, gypsiferous soils are often associated with distinct floras and high abdundances of biocrusts, especially lichens. At local to micro-scales, biocrusts often are better developed in habitats with lower radiation loads such as polar-oriented slopes, or shaded habitats. Also at small scales, vascular plant canopies buffer microclimate for biocrusts, but also exert negative influences such as burial by litter. While our knowledge of biocrust distribution has advanced rapidly, there are considerable geographic and taxonomic gaps in our knowledge and a pronounced lack of truly global studies.

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Patterns and controls on nitrogen cycling of biological soil crusts

Biocrusts play a significant role in the nitrogen [N ] cycle within arid and semi-arid ecosystems, as they contribute major N inputs via biological fixation and dust capture, harbor internal N transformation processes, and direct N losses via N dissolved, gaseous and erosional loss processes (Fig. 1). Because soil N availability in arid and semi-arid ecosystems is generally low and may limit net primary production (NPP), especially during periods when adequate water is available, understanding the mechanisms and controls of N input and loss pathways in biocrusts is critically important to our broader understanding of N cycling in dryland environments. In particular, N cycling by biocrusts likely regulates short-term soil N availability to support vascular plant growth, as well as long-term N accumulation and maintenance of soil fertility. In this chapter, we review the influence of biocrust nutrient input, internal cycling, and loss pathways across a range of biomes. We examine linkages between N fixation capabilities of biocrust organisms and spatio-temporal patterns of soil N availability that may influence the longer-term productivity of dryland ecosystems. Lastly, biocrust influence on N loss pathways such as N gas loss, leakage of N compounds from biocrusts, and transfer in wind and water erosion are important to understand the maintenance of dryland soil fertility over longer time scales. Although great strides have been made in understanding the influence of biocrusts on ecosystem N cycling, there are important knowledge gaps in our understanding of the influence of biocrusts on ecosystem N cycling that should be the focus of future studies. Because work on the interaction of N cycling and biocrusts was reviewed in Belnap and Lange (2003), this chapter will focus primarily on research findings that have emerged over the last 15 years (2000-2015).

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Carbon budgets of biological soil crusts at micro-, meso-, and global scales

The importance of biocrusts in the ecology of arid lands across all continents is widely recognized. In spite of this broad distribution, contributions of biocrusts to the global biogeochemical cycles have only recently been considered. While these studies opened a new view on the global role of biocrusts, they also clearly revealed the lack of data for many habitats and of overall standards for measurements and analysis. In order to understand carbon cycling in biocrusts and the progress which has been made during the last 15 years, we offer a multi-scale approach covering different climatic regions. We also include a discussion on available measurement techniques at each scale: A micro-scale section focuses on the individual organism level, including modeling based on the combination of field and lab data. The meso-scale section addresses the CO2 exchange of a complete ecosystem or at the community level. Finally, we consider the contribution of biocrusts at a global scale, giving a general perspective of the most relevant findings regarding the role of biological soil crusts in the global terrestrial carbon cycle.

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Biological soil crusts as soil stabilizers

Soil erosion is of particular concern in dryland regions, as the sparse cover of vascular plants results in large interspaces unprotected from the erosive forces of wind and water. Thus, most of these soil surfaces are stabilized by physical or biological soil crusts. However, as drylands are extensively used by humans and their animals, these crusts are often disturbed, compromising their stabilizing abilities. As a result, approximately 17.5% of the global terrestrial lands are currently being degraded by wind and water erosion. All components of biocrusts stabilize soils, including green algae, cyanobacteria, fungi, lichens, and bryophytes, and as the biomass of these organisms increases, so does soil stability. In addition, as lichens and bryophytes live atop the soil surface, they provide added protection from raindrop impact that cyanobacteria and fungi, living within the soil, cannot. Much research is still needed to determine the relative ability of individual species and suites of species to stabilize soils. We also need a better understanding of why some individuals or combination of species are better than others, especially as these organisms become more frequently used in restoration efforts.

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Biological soil crusts: An organizing principle in dryland ecosystems (aka: the role of biocrusts in arid land hydrology)

Biocrusts exert a strong influence on hydrological processes in drylands by modifying numerous soil properties that affect water retention and movement in soils. Yet, their role in these processes is not clearly understood due to the large number of factors that act simultaneously and can mask the biocrust effect. The influence of biocrusts on soil hydrology depends on biocrust intrinsic characteristics such as cover, composition, and external morphology, which differ greatly among climate regimes, but also on external factors as soil type, topography and vegetation distribution patterns, as well as interactions among these factors. This chapter reviews the most recent literature published on the role of biocrusts in infiltration and runoff, soil moisture, evaporation and non-rainfall water inputs (fog, dew, water absorption), in an attempt to elucidate the key factors that explain how biocrusts affect land hydrology. In addition to the crust type and site characteristics, recent studies point to the crucial importance of the type of rainfall and the spatial scale at which biocrust effects are analyzed to understand their role in hydrological processes. Future studies need to consider the temporal and spatial scale investigated to obtain more accurate generalizations on the role of biocrusts in land hydrology.

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How biological soil crusts became recognized as a functional unit: a selective history

It is surprising that despite the world-wide distribution and general importance of biological soil crusts (biocrusts), scientific recognition and functional analysis of these communities is a relatively young field of science. In this chapter, we sketch the historical lines that led to the recognition of biocrusts as a community with important ecosystem functions. The idea of biocrusts as a functional ecological community has come from two main scientific branches: botany and soil science. For centuries, botanists have long recognized that multiple organisms colonize the soil surface in the open and often dry areas occurring between vascular plants. Much later, after the initial taxonomic and phyto-sociological descriptions were made, soil scientists and agronomists observed that these surface organisms interacted with soils in ways that changed the soil structure. In the 1970’s, research on these communities as ecological units that played an important functional role in drylands began in earnest, and these studies have continued to this day. Here, we trace the history of these studies from the distant past until 1990, when biocrusts became well-known to scientists and the public.

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Biocrusts in the context of global change

A wide range of studies show global environmental change will profoundly affect the structure, function, and dynamics of terrestrial ecosystems. The research synthesized here underscores that biocrust communities are also likely to respond significantly to global change drivers, with a large potential for modification to their abundance, composition, and function. We examine how elevated atmospheric CO 2 concentrations, climate change (increased temperature and altered precipitation), and nitrogen deposition affect biocrusts and the ecosystems they inhabit. We integrate experimental and observational data, as well as physiological, community ecology, and biogeochemical perspectives. Taken together, these data highlight the potential for biocrust organisms to respond dramatically to environmental change and show how changes to biocrust community composition translate into effects on ecosystem function (e.g., carbon and nutrient cycling, soil stability, energy balance). Due to the importance of biocrusts in regulating dryland ecosystem processes and the potential for large modifications to biocrust communities, an improved understanding and predictive capacity regarding biocrust responses to environmental change are of scientific and societal relevance.

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Natural recovery of biological soil crusts after disturbance

Natural recovery of biological soil crusts (biocrusts) is influenced by a number of different parameters, such as climate, soil conditions, the severity of disturbance, and the timing of disturbance relative to the climatic conditions. In recent studies, it has been shown that recovery is often not linear, but a highly dynamic process directly influenced by non-linear external parameters as extraordinary climatic conditions (e.g., particularly dry or wet year). Natural recovery often follows a general succession pattern, starting out with cyanobacteria and algae, which is then followed by lichens and bryophytes at a later stage. However, this general sequence can be altered by parameters like dust deposition, fire effects, and special climatic conditions as in fog deserts and under mesic climates. Recent studies have proposed that under favorable, stable soil conditions, the initial soil-stabilizing cyanobacteria-dominated succession stages may be omitted and moss-dominated biocrusts can develop in the initial phases of biocrust development. During natural recovery of biocrusts, soil properties change, e.g., soil nutrient and organic matter contents increase. Also, silt and clay contents of encrusted soils increase with biocrust maturity, which may be caused by two mechanisms, i.e. entrapment of fine soil particles by biocrusts and the new formation of smaller particles by weathering of the existing substrate.

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