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Alternative pathways to landscape transformation: Invasive grasses, burn severity and fire frequency in arid ecosystems

Arid ecosystems are often vulnerable to transformation to invasive-dominated states following fire, but data on persistence of these states are sparse. The grass/fire cycle is a feedback process between invasive annual grasses and fire frequency that often leads to the formation of alternative vegetation states dominated by the invasive grasses. However, other components of fire regimes, such as burn severity, also have the potential to produce long-term vegetation transformations. Our goal was to evaluate the influence of both fire frequency and burn severity on the transformation of woody-dominated communities to communities dominated by invasive grasses in major elevation zones of the Mojave Desert of western North America. We used a chronosequence design to collect data on herbaceous and woody cover at 229 unburned reference plots and 578 plots that burned between 1972 and 2010. We stratified the plots by elevation zone (low, mid, high), fire frequency (1–3 times) and years post-fire (YPF; 1–5, 6–10, 11–20 and 21–40 YPF). Burn severity for each plot was estimated by the difference normalized burn ratio. We identified two broad post-fire successional pathways. One was an outcome of fire frequency, resulting in a strong potential transformation via the grass/fire cycle. The second pathway was driven by burn severity, the critical aspect being that long-term transformation of a community could occur from just one fire in areas that burned at high or sometimes moderate severity. Dominance by invasive grasses was most likely to occur in low-and high-elevation communities; cover of native herbaceous species was often greater than that of invasive grasses in the mid-elevation zone. Synthesis . Invasive grasses can dominate a site that burned only one time in many decades at high severity, or a site that burned at low severity but multiple times in the same time period. However, high burn severity may predispose areas to more frequent fire because they have relatively high cover of invasive annual grass, suggesting burn severity and fire frequency have both independent and synergistic effects. Resilience in vegetation structure following fire in many arid communities may be limited to a narrow window of low burn severity in areas that have not burned in many decades.

Mojave Desert

Selection of vegetation types and density of bison in an arid ecosystem

Understanding species habitat selection and factors that drive selection are key components for conservation. We report the first resource selection functions (RSFs) for bison inhabiting an arid ecosystem and use them with density estimates of bison to estimate the number of bison that could be supported if the bison range were expanded to federal lands in the San Luis Valley of Colorado. We derived RSFs for vegetation types using locations of plains bison collected weekly over 3 years from 2005 to 2007. Bison selected for wet or mesic grassland habitats in all seasons. Wetland selection by bison was predicted to be 18 times greater than that of rabbitbrush vegetation, the reference category, and selection of meadows was predicted to be 11 times greater than that of the rabbitbrush type. Willow-dominated plant communities were strongly avoided. Cottonwood communities were also avoided, with the exception of some moderate levels of selection in fall. The willow and cottonwood communities have an understory with low biomass of herbaceous species and low productivity in this arid system. Based on the RSFs we predicted that in the San Luis Valley of Colorado up to 2,379 bison could be supported in similar habitats under Fish and Wildlife Service (FWS) jurisdiction, and up to 759 bison could be supported on adjacent National Park Service (NPS) land. This modeling framework provides a conservation tool for the restoration of bison to their historical habitats, and has utility for application to other terrestrial species where assumptions are met.

Colorado

Effects of invasive plants on fire regimes and postfire vegetation diversity in an arid ecosystem

We assessed the impacts of co‐occurring invasive plant species on fire regimes and postfire native communities in the Mojave Desert, western USA. We analyzed the distribution and co‐occurrence patterns of three invasive annual grasses (Bromus rubens, Bromus tectorum, and Schismus spp.) known to alter fuel conditions and community structure, and an invasive forb (Erodium cicutarium) which dominates postfire sites. We developed species distribution models (SDMs) for each of the four taxa and analyzed field plot data to assess the relationship between invasives and fire frequency, years postfire, and the impacts on postfire native herbaceous diversity. Most of the Mojave Desert is highly suitable for at least one of the four invasive species, and 76% of the ecoregion is predicted to have high or very high suitability for the joint occurrence of B. rubens and B. tectorum and 42% high or very high suitability for the joint occurrence of the two Bromus species and E. cicutarium. Analysis of cover from plot data indicated two or more of the species occurred in 77% of the plots, with their cover doubling with each additional species. We found invasive cover in burned plots increased for the first 20 years postfire and recorded two to five times more cover in burned than unburned plots. Analysis also indicated that native species diversity and evenness as negatively associated with higher levels of relative cover of the four invasive taxa. Our findings revealed overlapping distributions of the four invasives; a strong relationship between the invasives and fire frequency; and significant negative impacts of invasives on native herbaceous diversity in the Mojave. This suggests predicting the distributions of co‐occurring invasive species, especially transformer species, will provide a better understanding of where native‐dominated communities are most vulnerable to transformations following fire or other disturbances.

California, Nevada

Biological soil crust and disturbance controls on surface hydrology in a semi-arid ecosystem

Biological soil crust communities (biocrusts) play an important role in surface hydrologic processes in dryland ecosystems, and these processes may then be dramatically altered with soil surface disturbance. In this study, we examined biocrust hydrologic responses to disturbance at different developmental stages on sandy soils on the Colorado Plateau. Our results showed that all disturbance (trampling, scalping and trampling+scalping) of the early successional light cyanobacterial biocrusts generally reduced runoff. In contrast, trampling well-developed dark-cyano-lichen biocrusts increased runoff and sediment loss relative to intact controls. Scalping did not increase runoff, implying that soil aggregate structure was important to the infiltration process. Well-developed, intact dark biocrusts generally had lower runoff, low sediment loss, and highest aggregate stability whereas the less-developed light biocrusts were highest in runoff and sediment loss when compared to the controls. These results suggest the importance of maintaining the well-developed dark biocrusts, as they are beneficial for lowering runoff and reducing soil loss and redistribution on the landscape. These data also suggest that upslope patches of light biocrust may either support water transport to downslope vegetation patches or alternatively this runoff may place dark biocrust patches at risk of disruption and loss, given that light patches increase runoff and thus soil erosion potential.

Colorado

Wildfire and the ecological niche: Diminishing habitat suitability for an indicator species within semi‐arid ecosystems

Globally accelerating frequency and extent of wildfire threatens the persistence of specialist wildlife species through direct loss of habitat and indirect facilitation of exotic invasive species. Habitat specialists may be especially prone to rapidly changing environmental conditions because their ability to adapt lags behind the rate of habitat alteration. As a result, these populations may become increasingly susceptible to ecological traps by returning to suboptimal breeding habitats that were dramatically altered by disturbance. We demonstrate a multistage modeling approach that integrates habitat selection and survival during the key nesting life‐stage of a bird species of high conservation concern, the greater sage‐grouse ( Centrocercus urophasianus ; hereafter, sage‐grouse). We applied these spatially explicit models to a spatiotemporally robust dataset of sage‐grouse nest locations and fates across wildfire‐altered sagebrush ecosystems of the Great Basin ecoregion, western United States. Female sage‐grouse exhibited intricate habitat selection patterns that varied across regional gradients of ecological productivity among sagebrush communities, but often selected nest sites that disproportionately resulted in nest failure. For example, 23% of nests occurred in wildfire‐affected habitats characterized by reduced sagebrush cover and greater composition of invasive annual grasses. We found survival of nests was negatively associated with wildfire‐affected areas, but positively associated with higher elevations with increased ruggedness and overall shrub cover. Strong site fidelity likely drove sage‐grouse to continue nesting in habitats degraded by wildfire. Hence, increasing frequency and extent of wildfire may contribute disproportionately to reduced reproductive success by creating ecological traps that act as population sinks. Identifying such habitat mismatches between selection and survival facilitates deeper understanding of the mechanisms driving reduced geographic niche space and population decline at broad spatiotemporal scales, while guiding management actions to areas that would be most beneficial to the species.

Global Change Biology

Hyperspectral remote sensing tools for quantifying plant litter and invasive species in arid ecosystems

Green vegetation can be monitored and distinguished using visible and infrared multiband and hyperspectral remote sensing methods. The problem has been in identifying and distinguishing the nonphotosynthetically active radiation (PAR) landscape components, such as litters and soils, from green vegetation [35-38]. Additionally, distinguishing different species of green vegetation is challenging using the relatively few bands available on most satellite sensors. This chapter focuses both on previously published work by Nagler et al. [35-38] that identified hyperspectral remote sensing characteristics that distinguish between green vegetation, soil, and litter (or senescent vegetation), and on new research conducted to aid in distinguishing invasive species from the mixed landcover surface.

Book chapter

Spatial estimates of soil moisture for understanding ecological potential and risk: a case study for arid and semi-arid ecosystems

Soil temperature and moisture (soil-climate) affect plant growth and microbial metabolism, providing a mechanistic link between climate and growing conditions. However, spatially explicit soil-climate estimates that can inform management and research are lacking. We developed a framework to estimate spatiotemporal-varying soil moisture (monthly, annual, and seasonal) and temperature-moisture regimes as gridded surfaces by enhancing the Newhall simulation model. Importantly, our approach allows for the substitution of data and parameters, such as climate, snowmelt, soil properties, alternative potential evapotranspiration equations and air-soil temperature offsets. We applied the model across the western United States using monthly climate averages (1981–2010). The resulting data are intended to help improve conservation and habitat management, including but not limited to increasing the understanding of vegetation patterns (restoration effectiveness), the spread of invasive species and wildfire risk. The demonstrated modeled results had significant correlations with vegetation patterns—for example, soil moisture variables predicted sagebrush (R 2 = 0.51), annual herbaceous plant cover (R 2 = 0.687), exposed soil (R 2 = 0.656) and fire occurrence (R 2 = 0.343). Using our framework, we have the flexibility to assess dynamic climate conditions (historical, contemporary or projected) that could improve the knowledge of changing spatiotemporal biotic patterns and be applied to other geographic regions.

Land

Ecology of bison, elk, and vegetation in an arid ecosystem

Herbivory has profound effects on vegetation production and structure in many different plant communities. The influence of herbivory on plants and ultimately ecosystem processes is shaped by the types of plants consumed, the intensity of herbivory, the evolutionary history of grazing, and the availability of water and nutrients to plants. The effect of ungulate herbivores on vegetation is of great interest to ecologists, land managers and agriculturalists. In addition, the Department of Interior recently established a Bison Conservation Initiative to provide for the conservation and restoration of North American plains- and wood bison, which includes establishing new populations and expanding existing populations. The San Luis Valley, Colorado, is being considered as a potential location for a bison conservation herd. Resource managers need to know the vegetation impacts of adding a second large ungulate to a system that already has elk.

Colorado

Food habits of rodents inhabiting arid and semi-arid ecosystems of central New Mexico

In this study, we describe seasonal dietary composition for 15 species of rodents collected in all major habitats on the Sevilleta National Wildlife Refuge (Socorro County) in central New Mexico. A comprehensive literature review of food habits for these species from throughout their distribution also is provided. We collected rodents in the field during winter, spring and late summer in 1998 from six communities: riparian cottonwood forest; piñon-juniper woodland; juniper-oak savanna; mesquite savanna; short-grass steppe; and Chihuahuan Desert scrubland. Rodents included Spermophilus spilosoma (Spotted Ground Squirrel), Perognathus flavescens (Plains Pocket Mouse), Perognathus flavus (Silky Pocket Mouse), Dipodomys merriami (Merriam’s Kangaroo Rat), Dipodomys ordii (Ord’s Kangaroo Rat), Dipodomys spectabilis (Banner-tailed Kangaroo Rat), Reithrodontomys megalotis (Western Harvest Mouse), Peromyscus boylii (Brush Mouse), Peromyscus eremicus (Cactus Mouse), Peromyscus leucopus (White-footed Mouse), Peromyscus truei (Piñon Mouse), Onychomys arenicola (Mearn’s Grasshopper Mouse), Onychomys leucogaster (Northern Grasshopper Mouse), Neotoma albigula/leucodon (White-throated Woodrats), and Neotoma micropus (Southern Plains Woodrat). We collected stomach contents of all species, and cheek-pouch contents of heteromyids, and quantified them in the laboratory. We determined seasonal diets in each habitat by calculating mean percentage volumes of seeds, arthropods and green vegetation (plant leaves and stems) for each species of rodent. Seeds consumed by each rodent were identified to genus, and often species, and quantified by frequency counts. Comparisons of diets between and among species of rodents, seasons, and ecosystems were also examined. We provide an appendix of all plant taxa documented.

Special publication of the Museum of Southwestern

Climatic controls on soil carbon accumulation and loss in a dryland ecosystems

Arid and semiarid ecosystems drive year-to-year variability in the strength of the terrestrial carbon (C) sink, yet there is uncertainty about how soil C gains and losses contribute to this variation. To address this knowledge gap, we embedded C-depleted soil mesocosms, containing litter or biocrust C inputs, within an in situ dryland ecosystem warming experiment. Over the course of one year, changes in microbial biomass and total soil organic C pools were monitored alongside hourly measurements of soil CO 2 flux. We also developed a biogeochemical model to explore the mechanisms that gave rise to observed soil C dynamics. Field data and model simulations demonstrated that water exerted much stronger control on soil biogeochemistry than temperature, with precipitation events triggering large CO 2 pulses and transport of litter- and biocrust-derived C into the soil profile. We expected leaching of organic matter would result in steady accumulation of C within the mineral soil over time. Instead, the size of the total organic C pool fluctuated throughout the year, largely in response to microbial growth: increases in the size of microbial biomass were negatively correlated with the quantity of C residing in the top 2 cm, where most biogeochemical changes were observed. Our data and models suggest that microbial responses to precipitation events trigger rapid metabolism of dissolved organic C inputs, which strongly limit accumulation of autotroph-derived C belowground. Accordingly, changes in the magnitude and/or frequency of precipitation events in this dryland ecosystem could have profound impacts on the strength of the soil C sink.

Utah

A multiscale, hierarchical model of pulse dynamics in arid-land ecosystems

Ecological processes in arid lands are often described by the pulse-reserve paradigm, in which rain events drive biological activity until moisture is depleted, leaving a reserve. This paradigm is frequently applied to processes stimulated by one or a few precipitation events within a growing season. Here we expand the original framework in time and space and include other pulses that interact with rainfall. This new hierarchical pulse-dynamics framework integrates space and time through pulse-driven exchanges, interactions, transitions, and transfers that occur across individual to multiple pulses extending from micro to watershed scales. Climate change will likely alter the size, frequency, and intensity of precipitation pulses in the future, and arid-land ecosystems are known to be highly sensitive to climate variability. Thus, a more comprehensive understanding of arid-land pulse dynamics is needed to determine how these ecosystems will respond to, and be shaped by, increased climate variability.

Annual Review of Ecology, Evolution, and Systemati

Understanding the effect of fire on vegetation composition and gross primary production in a semi-arid shrubland ecosystem using the Ecosystem Demography (EDv2.2) model

Wildfires in sagebrush (Artemisia spp.) dominated semi-arid ecosystems in the western United States have increased dramatically in frequency and severity in the last few decades. Severe wildfires often lead to the loss of native sagebrush communities and change the biogeochemical conditions which make it difficult for sagebrush to regenerate. Invasion of cheat- grass (Bromus tectorum) accentuates the problem by making the ecosystem more susceptible to frequent burns. Managers have implemented several techniques to cope with the cheatgrass-fire cycle, ranging from controlling undesirable fire effects by removing fuel loads either mechanically or via prescribed burns, to seeding the fire-affected areas with shrubs and native perennial forbs. There have been a number of studies at local scales to understand the direct impacts of wildfire on vegetation, however, there is a larger gap in understanding these impacts at broad spatial and temporal scales. This need highlights the importance of dynamic global vegetation models (DGVMs) and remote sensing. In this study, we explored the influence of fir on vegetation composition and gross primary production (GPP) in the sagebrush ecosystem using the Ecosystem Demography (EDv2.2) model, a dynamic global vegetation model. We selected Reynolds Creek Experimental Watershed (RCEW) to run our simulation study, an intensively monitored sagebrush-dominated ecosystem in the northern Great Basin. We ran point-based simulations at four existing flux-tower sites in the study area for a total 150 years after turning on the fire module in the 25th year. Results suggest dominance of shrubs in a non-fire scenario, however under the fire scenario we observed contrasting phases of high and low shrub density and C3 grass growth. Regional model simulations showed a gradual decline in GPP for fire-introduced areas through the initial couple of years instead of killing all the vegetation in the affected area in the first year itself. We also compared the results from EDv2.2 with satellite-derived GPP estimates for the areas in RCEW burned by a wildfire in 2015 (Soda Fire). We observed moderate pixel-level correlations between maps of post-fire recovery EDv2.2 GPP and MODIS derived GPP. This study contributes to understanding the application of ecosystem models to investigate temporal dynamics of vegetation under alternative fire regimes and post-fire ecosystem restoration.

Idaho

Fire frequency impacts soil properties and processes in sagebrush steppe ecosystems of the Columbia Basin

Increased fire frequency in semi-arid ecosystems can alter biochemical soil properties and soil processes that underpin ecosystem structure and functioning, thus threatening native plant communities and the species that rely on them. However, there is much uncertainty about the magnitude of change as soils are exposed to more fires, because soil recovery and changes in fire severity following a first fire mediate the impact of successive fires on soil properties. With this study we aim to evaluate how increased fire frequency affects soil biochemical properties (i.e. soil pH, soil organic matter (SOM), soil organic carbon (SOC), soil structure and mineral N) and processes (i.e. microbial and enzymatic activity) in a sagebrush-steppe ecosystem located in the Columbia Plateau Ecoregion, Washington, USA. During 2016, we collected soils from once (2012), twice (2003 and 2012), and thrice (2003, 2007, and 2012) burned areas, enabling us to test the hypothesis that increasing fire frequency will exacerbate the impact of fire on soil properties and processes. Our study yielded three main results: (1) fire reduced the total soil C concentration and soil C in aggregates relative to unburned soil, but only when soil was exposed to fire once (i.e. the most recent fire), (2) compared to the unburned soils, SOM contents, enzyme activity and microbial CO 2 respiration were suppressed in the once and thrice burned soils, but not in the twice burned soils, and (3) fire increased NO 3 − -N contents across the once and twice burned sites, and reduced enzyme activity associated with N cycling in the thrice burned sites. Taken together, our findings suggest that a one-time fire in this shrub dominated semi-arid ecosystem significantly changes soil biochemical attributes and microbially driven processes. With sufficient time between fires, these structural and functional properties can partially recover, and this may persist even after a second fire, but recovery is limited when a third fire creates an additional disturbance at a shorter time interval. Furthermore, while soil C pools and microbial decomposition processes were able to recover with sufficient time, greater soil resource availability prevailed in soil across all fire frequencies, indicating that fire is likely to promote invasion and reduce ecosystem stability, even when other soil properties recover.

Washington

Management implications of the ecology of free-roaming horses in semiarid ecosystems of the western United States

Compared to other ungulates of North America, free - roaming horses ( Equus caballus ) possess a unique evolutionary history that has given rise to a distinct suite of behavioral, morphological, and physiological traits. Because of their unique combination of cecal digestion, an elongate head with flexible lips, and non-uniform use of the landscape, horses represent a unique disturbance agent in semi-arid ecosystems of the western United States . Consequently, it is inappropriate to assume that influences of horses on the structure, composition, function, and pattern of arid and semi-arid ecosystems will mirror influences of cattle or other artiodactyls. Although management areas for free - roaming horses occupy 18.6 million ha of land across western North America, we know relatively little about how western ecosystems and their components have responded to this uniquely managed ungulate. I draw on my research of horse habitats in the western Great Basin (U.S.A.) to examine predictions of horses ' unique influence, and advocate for continued research to refine our understanding of synecological relationships among horses and diverse ecosystem components in arid and semi-arid regions.

Wildlife Society Bulletin

Mercury cycling across a U.S. semi-arid mountain ecosystem elevation gradient

Mountains comprise ∼30% of the Earth's surface, but mercury (Hg) cycling in these regions remains understudied, particularly in the semi-arid western U.S. where strong climatic and ecological gradients in mountainous landscapes influence Hg deposition, retention, and bioaccumulation. In this study, we quantified growing season inputs, storage, and bioaccumulation of Hg along a ∼2,000 m elevation gradient in the Colorado Rocky Mountains, spanning the plains to the alpine. We measured Hg in atmospheric deposition, vegetation, soil, and 12-day-old chickadees. Accounting for percent canopy cover, open precipitation was the largest component of atmospheric deposition at all elevations, followed by throughfall and litterfall fluxes. Atmospheric Hg fluxes peaked at mid-elevations, likely due to cloud-cap dynamics and denser canopy cover. Total gaseous Hg and precipitation fluxes were highest at low elevations, likely reflecting local emissions and meteorological pooling. Surface soil Hg storage was more strongly predicted by organic matter content ( R 2 = 0.49; p < 0.01) and water retention ( R 2 = 0.45; p < 0.01) than by elevation ( R 2 = 0.21; p < 0.05). Alpine soils (66.3 ± 25.3 ng g −1 ) had significantly higher total Hg concentrations than lower elevations (<41.0 ± 12.7 ng g −1 ; p < 0.01), likely reflecting slower organic matter turnover. Soils on north-facing slopes also retained significantly higher pools of Hg in surface soils compared with south- and east-facing slopes. Vegetation Hg pools were greatest in the alpine region, likely due to long-lived plant species. Methylmercury (MeHg) concentrations in chickadee feathers peaked at mid-elevations (205 ± 155 ng g −1 ), corresponding to higher ecosystem Hg inputs via throughfall. Our results show that deposition, canopy cover, and meteorological conditions—not elevation alone—predict Hg retention and bioaccumulation.

Colorado

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).

Book chapter

Soil respiration patterns and controls in limestone cedar glades

Aims Drivers of soil respiration ( R s ) in rock outcrop ecosystems remain poorly understood. We investigated these drivers in limestone cedar glades, known for their concentrations of endemic plant species and for seasonal hydrologic extremes (xeric and saturated conditions), and compared our findings to those in temperate grasslands and semi-arid ecosystems. Methods We measured R s , soil temperature ( T s ), volumetric soil water content (SWC), soil organic matter (SOM), soil depth, and vegetation cover monthly over 16 mo and analyzed effects of these variables on R s . Results Seasonally, R s primarily tracked T s (r 2 =0.77; P < 0.01); however R s was depressed during a summer drought. SOM was highly variable spatially, and incorporating SOM effects into the R s model dramativally improved model performance. Both shallow soil and sparse vegetation cover were also associated with lower R s . Conclusions Soil depth, SOM, and vegetation cover were important drivers of R s in limestone cedar glades. Seasonal R s patterns reflected those for mesic temperate grasslands more than for semi-arid ecosystems, in that R s primarily tracked temperature for most of the year.

Plant and Soil