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Bioclimatic envelopes for individual demographic events driven by extremes: Plant mortality from drought and warming

The occurrence of plant species across the globe is largely constrained by climate. Ecologists use plant-climate relationships such as bioclimatic envelopes and related niche models to determine potential environmental conditions promoting probable species occurrence. Traditionally bioclimatic envelopes either exclude disturbance explicitly, or only include disturbance as infrequent and smaller scale processes, assuming that the net effect of climate parameters on key demographic processes predict longer-term equilibrial responses of biota. Due to increasing frequency and extent of extreme events associated with climate change, ecologists may need to increase focus on individual demographic events driven by environmental extremes such as widespread coral bleaching or large-scale tree die-off. An expanded focus on how extreme events catalyze individual demographic events would complement existing tools that predict long-term equilibrial biogeographic responses associated with long-term trends in climate. In many cases, extreme conditions (e.g. drought) are a necessary precursor for an abrupt demographic event (e.g. large-scale tree die-off) and the effects of extremes can be exacerbated by climatic trends (e.g. higher temperatures in combination with drought). Here, we highlight application of bioclimatic models for predicting individual demographic events. Defining the environmental conditions that precipitate demographic events such as widespread tree mortality is a necessary precursor for applying predictions to geographic space, and may require challenging biota with experiments that impose a combination of ecologically extreme conditions in one parameter and a shifting distribution in another (e.g. drought under higher temperatures). Currently data on conditions that drive individual demographic events associated with extremes are usually rare, aggregated across time, and/or correlative. We highlight this approach with a case study of drought-induced mortality in adult Pinus edulis trees that predicts a more than five-fold increase in frequency of die-off events under a global change scenario of high emissions. This general approach complements both traditional bioclimatic envelopes and more detailed physiological approaches currently being refined to address climate change challenges. Notably, this proposed approach could be developed for any climate condition or plant life stage, offering promise for improving predictions of individual demographic events that are rapidly altering ecosystems globally.

International Journal of Plant Sciences

Root growth and function of three Mojave Desert grasses in response to elevated atmospheric CO2 concentration

Root growth and physiological responses to elevated CO 2 were investigated for three important Mojave Desert grasses: the C 3 perennial Achnatherum hymenoides , the C 4 perennial Pleuraphis rigida and the C 3 annual Bromus madritensis ssp. rubens . Seeds of each species were grown at ambient (360 μl l −1 ) or elevated (1000 μl l −1 ) CO 2 in a glasshouse and harvested at three phenological stages: vegetative, anthesis and seed fill. Because P. rigida did not flower during the course of this study, harvests for this species represent three vegetative stages. Primary productivity was increased in both C 3 grasses in response to elevated CO 2 (40 and 19% for A. hymenoides and B. rubens , respectively), but root biomass increased only in the C 3 perennial grass. Neither above-ground nor below-ground biomass of the C 4 perennial grass was significantly affected by the CO 2 treatment. Elevated CO 2 did not significantly affect root surface area for any species. Total plant nitrogen was also not statistically different between CO 2 treatments for any species, indicating no enhanced uptake of N under elevated CO 2 . Physiological uptake capacities for NO 3 and NH 4 were not affected by the CO 2 treatment during the second harvest; measurements were not made for the first harvest. However, at the third harvest uptake capacity was significantly decreased in response to elevated CO 2 for at least one N form in each species. NO 3 uptake rates were lower in A. hymenoides and P. rigida , and NH 4 uptake rates were lower in B. rubens at elevated CO 2 . Nitrogen uptake on a whole root-system basis (NO 3 +NH 4 uptake capacity × root biomass) was influenced positively by elevated CO 2 only for A. hymenoides after anthesis. These results suggest that elevated CO 2 may result in a competitive advantage for A. hymenoides relative to species that do not increase root-system N uptake capacity. Root respiration measurements normalized to 20 °C were not significantly affected by the CO 2 treatment. However, specific root respiration was significantly correlated with either root C∶N ratio or root water content when all data per species were included within a simple regression model. The results of this study provide little evidence for up-regulation of root physiology in response to elevated CO 2 and indicate that root biomass responses to CO 2 are species-specific.

New Phytologist

Tropicalization of temperate ecosystems in North America: The northward range expansion of tropical organisms in response to warming winter temperatures

Tropicalization is a term used to describe the transformation of temperate ecosystems by poleward‐moving tropical organisms in response to warming temperatures. In North America, decreases in the frequency and intensity of extreme winter cold events are expected to allow the poleward range expansion of many cold‐sensitive tropical organisms, sometimes at the expense of temperate organisms. Although ecologists have long noted the critical ecological role of winter cold temperature extremes in tropical–temperate transition zones, the ecological effects of extreme cold events have been understudied, and the influence of warming winter temperatures has too often been left out of climate change vulnerability assessments. Here, we examine the influence of extreme cold events on the northward range limits of a diverse group of tropical organisms, including terrestrial plants, coastal wetland plants, coastal fishes, sea turtles, terrestrial reptiles, amphibians, manatees, and insects. For these organisms, extreme cold events can lead to major physiological damage or landscape‐scale mass mortality. Conversely, the absence of extreme cold events can foster population growth, range expansion, and ecological regime shifts. We discuss the effects of warming winters on species and ecosystems in tropical–temperate transition zones. In the 21st century, climate change‐induced decreases in the frequency and intensity of extreme cold events are expected to facilitate the poleward range expansion of many tropical species. Our review highlights critical knowledge gaps for advancing understanding of the ecological implications of the tropicalization of temperate ecosystems in North America.

Global Change Biology

RIP-ET: A riparian evapotranspiration package for MODFLOW-2005

A new evapotranspiration package for the U.S. Geological Survey's groundwater-flow model, MODFLOW, is documented. The Riparian Evapotranspiration Package (RIP-ET) provides flexibility in simulating riparian and wetland transpiration not provided by the Evapotranspiration (EVT) or Segmented Function Evapotranspiration (ETS1) Packages for MODFLOW 2005. This report describes how the RIP-ET package was conceptualized and provides input instructions, listings and explanations of the source code, and an example. Traditional approaches to modeling evapotranspiration (ET) processes assume a piecewise linear relationship between ET flux and hydraulic head. The RIP-ET replaces this traditional relationship with a segmented, nonlinear dimensionless curve that reflects the eco-physiology of riparian and wetland ecosystems. Evapotranspiration losses from these ecosystems are dependent not only on hydraulic head, but on the plant types present. User-defined plant functional groups (PFGs) are used to elucidate the interaction between plant transpiration and groundwater conditions. Five generalized plant functional groups based on transpiration rates, plant rooting depth, and water tolerance ranges are presented: obligate wetland, shallow-rooted riparian, deep-rooted riparian, transitional riparian and bare ground/open water. Plant functional groups can be further divided into subgroups (PFSGs) based on plant size, density or other characteristics. The RIP-ET allows for partial habitat coverage and mixtures of plant functional subgroups to be present in a single model cell. RIP-ET also distinguishes between plant transpiration and bare-ground evaporation. Habitat areas are designated by polygons; each polygon can contain a mixture of PFSGs and bare ground, and is assigned a surface elevation. This process requires a determination of fractional coverage for each of the plant functional subgroups present in a polygon to account for the mixture of coverage types and resulting transpiration. The fractional cover within a cell has two components: (1) the polygonal fraction of active habitat (excluding area of bare ground, dead trees, or brush) in a cell, and (2) fraction of plant type area or bare ground area in a polygon. RIP-ET determines the transpiration rate for each plant functional group and evaporation from bare ground/open water in a cell, the total ET in the cell, and the total ET rate over the region of simulation.

Techniques and Methods

Habitat and sex differences in physiological condition of breeding Southwestern Willow Flycatchers ( Empidonax traillii extimus )

The Southwestern Willow Flycatcher ( Empidonax traillii extimus ; here- after “flycatcher”) is a federally listed endangered species that breeds in densely vegetated riparian habitats dominated by native and exotic plants, including introduced monotypic saltcedar (Tamarix ramosissima) . Some workers have theorized that saltcedar is unsuitable habitat for the flycatcher, primarily because it generally supports a smaller and less diverse invertebrate community (the flycatcher's food base) than native habitats (e.g. Salix spp.). However, differences in insect communities between native and saltcedar habitats are not proof that saltcedar habitats are inferior. The only way to evaluate whether the habitats differ in dietary or energetic quality is to document actual food limitation or its manifestations. Measurements of an individual's body condition and metabolic state can serve as indicators of environmental stressors, such as food limitation and environmental extremes. We captured 130 flycatchers breeding in native and saltcedar habitats in Arizona and New Mexico and measured 12 variables of physiological condition. These variables included body mass, fat level, body condition index, hematocrit, plasma triglycerides, plasma free fatty acids and glycerol, plasma glucose and beta-hydroxybutyrate, plasma uric acid, total leukocyte count, and heterophil-to-lymphocyte ratio. We found substantial sex-based differences in the condition of male and female flycatchers. Ten of the 12 measures of physiological condition differed significantly between the sexes. In all cases where male and female condition differed (except mass), the differences suggest that males were in poorer condition than females. We found few habitat-based differences in flycatcher condition. Only 3 of the 12 physiological condition indices differed significantly between habitats. Our data show that, at least in some parts of the flycatcher's range, there is no evidence that flycatchers breeding in saltcedar habitats exhibit poorer nutritional condition or are suffering negative physiological affects.

The Auk

Tidal freshwater forested wetlands: Future research needs and an overview of restoration: Chapter 17

Studies of tidal freshwater forested wetlands are few in contrast to the diversity of conditions and information needs that exist for this ecosystem type. Basic information is lacking on the physiological ecology of major wetland tree species under natural settings, the structure and dynamics of pure and mixed species communities, soil-plant interactions, biogeochemistry, hydrology, soils, wildlife habitat, primary biotic and abiotic functions, and the response of these systems to natural and human-caused disruptions. Existing information is often not in a form that can be applied to ecosystem problems, especially those related to management, restoration, or creation of tidal swamps. Accordingly, there is a critical need for research on fundamental biotic and abiotic processes and functions in tidal forested wetland landscapes on a local and regional scale. In this chapter, we detail those research needs, and we highlight some restoration ideas for tidal freshwater forested wetlands with the hope that much additional research will follow.

Book chapter

**1**5N-NMR INVESTIGATION OF HYDROXYLAMINE DERIVATIZED HUMIC SUBSTANCES.

Humic substances are the most abundant naturally occurring refactory organic compounds in soils and water. They have a broad range of physical, chemical and physiological properties. In soils, humic substances contribute to the cation exchange capacity, help maintain the physical structure, and play a role in plant growth and nutrition. In aquatic systems, humic substances serve to regulate the levels of inorganic constituents, yield trihalomethanes upon chlorination, and transport or concentrate organic and inorganic pollutants. The oxygen containing functional groups of humic and fulvic acids are believed to play a key role in the chemical properties of humic substances. This study was undertaken to gain additional information on the specific types of oxygen functionalities in humic substances. Since the analysis of hydroxyl moieties had been earlier established, we focused our attention on the analysis of ketone and aldehyde functional groups in humic substances.

Conference Paper

Characterizing canopy biochemistry from imaging spectroscopy and its application to ecosystem studies

For two decades, remotely sensed data from imaging spectrometers have been used to estimate non-pigment biochemical constituents of vegetation, including water, nitrogen, cellulose, and lignin. This interest has been motivated by the important role that these substances play in physiological processes such as photosynthesis, their relationships with ecosystem processes such as litter decomposition and nutrient cycling, and their use in identifying key plant species and functional groups. This paper reviews three areas of research to improve the application of imaging spectrometers to quantify non-pigment biochemical constituents of plants. First, we examine recent empirical and modeling studies that have advanced our understanding of leaf and canopy reflectance spectra in relation to plant biochemistry. Next, we present recent examples of how spectroscopic remote sensing methods are applied to characterize vegetation canopies, communities and ecosystems. Third, we highlight the latest developments in using imaging spectrometer data to quantify net primary production (NPP) over large geographic areas. Finally, we discuss the major challenges in quantifying non-pigment biochemical constituents of plant canopies from remotely sensed spectra.

Remote Sensing of Environment

Effects and empirical critical loads of Nitrogen for ecoregions of the United States

Human activity in the last century has increased nitrogen (N) deposition to a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. We synthesized current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and estimated associated empirical critical loads of N for several receptors: freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Biogeochemical responses included increased N mineralization and nitrification, increased gaseous N losses, and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root-shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species. The range of critical loads of nutrient N reported for U.S. ecoregions, inland surface waters, and freshwater wetlands is 1–39 kg N ha−1 yr−1, spanning the range of N deposition observed over most of the country. The empirical critical loads of N tend to increase in the following sequence: diatoms, lichens and bryophytes, mycorrhizal fungi, herbaceous plants and shrubs, trees.

Book chapter

Climate adaption and post-fire restoration of a foundational perennial in cold desert: Insights from intraspecific variation in response to weather

1.The loss of foundational but fire-intolerant perennials such as sagebrush due to increases in fire size and frequency in semiarid regions has motivated efforts to restore them, often with mixed or even no success. Seeds of sagebrush Artemisia tridentata and related species must be moved considerable distances from seed source to planting sites, but such transfers have not been guided by an understanding of local climate adaptation. Initial seedling establishment and its response to weather are a key demographic bottleneck that likely varies among subspecies and populations of sagebrush. 2.We assessed differences in survival, growth, and physiological responses of sagebrush to weather among eleven seed sources that varied in subspecies, cytotype, and climates-of-origin over 18 months following outplanting. Diploid or polyploid populations of mountain, Wyoming, and basin big sagebrush ( A.tridentata ssp. vaseyana, A.tridentata ssp. wyomingensis, and A.tridentata ssp. tridentata , respectively) were planted onto five burned sites that normally support A.t.wyomingensis with some A.t.tridentata . 3.A.t.wyomingensis had the most growth and survival, and tetraploid populations had greater survival and height than diploids. Seasonal timing of mortality varied among the subspecies/cytotypes and was more closely related to minimum temperatures than water deficit. 4.Temperatures required to induce ice formation were up to 6°C more negative in 4n- A.t.tridentata and A.t.wyomingensis than other subspecies/cytotypes, indicating greater freezing avoidance. In contrast, freezing resistance of photosynthesis varied only 1°C among subspecies/cytotypes, being greatest in A.t.wyomingensis and least in the subspecies normally considered most cold-adapted, A.t.vaseyana . A large spectrum of reliance on freezing-avoidance vs. freezing-tolerance was observed and corresponded to differences in post-fire survivorship among subspecies/cytotypes. Differences in water deficit responses among subspecies/cytotypes were not as strong and did not relate to survival patterns. 5. Synthesis and applications. Low temperature responses are a key axis defining climate adaptation in young sagebrush seedlings and vary more with cytotype than with subspecies, which contrasts with the traditional emphases on (i) water limitations to explain establishment in these deserts, and (ii) subspecies in selecting restoration seedings. These important and novel insights on climate adaptation are critical for seed selection and parameterizing seed transfer zones, and were made possible by incorporating weather data with survival statistics. The survival/weather statistics used here could be applied to any restoration planting or seeding to help elucidate factors contributing to success and enable adaptive management.

Journal of Applied Ecology

Photosynthesis and fluctuating asymmetry as indicators of plant response to soil disturbance in the Fall-Line Sandhills of Georgia: a case study using Rhus copallinum and Ipomoea pandurata

We examined net photosynthesis, transpiration, stomatal conductance, and leaf fluctuating asymmetry on two species ( Rhus copallinum and Ipomoea pandurata ) as indicators of stress at nine sites across a gradient of soil disturbance at Fort Benning, Georgia. There were three sites for each of three disturbance levels. Physical habitat disturbance was caused by activities associated with infantry training, including mechanized elements (tanks and personnel carriers) and foot soldiers. In addition, we examined the influence of prescribed burns and microhabitat effects (within meter‐square quadrats centered about the plant) on these measures of plant stress. Net photosynthesis declined with increasing disturbance in the absence of burning for both species. However, when sites were burned the previous year, net photosynthesis increased with increasing disturbance. Developmental instability in Rhus , as measured by fluctuating asymmetry, also declined with increasing disturbance in the absence of burning but increased with disturbance if sites were burned the previous year. Developmental instability was much less sensitive to burning in Ipomoea and in general was lowest at intermediate disturbance sites. Microenvironmental and microhabitat effects were weakly correlated with measures of plant stress when all sites were combined. However, higher correlations were obtained within site categories, especially when the recent history of prescribed burning was used as a category. Finally, using all of the combined data in a discriminant function analysis allowed us to correctly predict the disturbance level of more than 80% of the plants. Plant stress is responsive to both large‐scale perturbations, such as burning, and microhabitat parameters. Because of this, it is important to include macro‐ and microhabitat parameters when assessing stress. Similarly, we found a combination of developmental and physiological indicators of stress was superior to using them separately.

International Journal of Plant Sciences

Climate change perils for dioecious plant species

Climate change, particularly increased aridity, poses a significant threat to plants and the biotic communities they support. Dioecious species may be especially vulnerable to climate change given that they often exhibit spatial segregation of the sexes, reinforced by physiological and morphological specialization of each sex to different microhabitats. In dimorphic species, the overexpression of a trait by one gender versus the other may become suppressed in future climates. Data suggest that males will generally be less sensitive to increased aridity than co-occurring females and, consequently, extreme male-biased sex ratios are possible in a significant number of populations. The effects of male-biased sex ratios are likely to cascade to dependent community members, especially those that are specialized on one sex.

Nature Plants

Pulse-drought atop press-drought: unexpected plant responses and implications for dryland ecosystems

In drylands, climate change is predicted to cause chronic reductions in water availability (press-droughts) through reduced precipitation and increased temperatures as well as increase the frequency and intensity of short-term extreme droughts (pulse-droughts). These changes in precipitation patterns may have profound ecosystem effects, depending on the sensitivities of the dominant plant functional types (PFTs). Here we present the responses of four Colorado Plateau PFTs to an experimentally imposed, 4-year, press-drought during which a natural pulse-drought occurred. Our objectives were to (1) identify the drought sensitivities of the PFTs, (2) assess the additive effects of the press- and pulse-drought, and (3) examine the interactive effects of soils and drought. Our results revealed that the C 3 grasses were the most sensitive PFT to drought, the C 3 shrubs were the most resistant, and the C 4 grasses and shrubs had intermediate drought sensitivities. Although we expected the C 3 grasses would have the greatest response to drought, the higher resistance of C 3 shrubs relative to the C 4 shrubs was contrary to our predictions based on the higher water use efficiency of C 4 photosynthesis. Also, the additive effects of press- and pulse-droughts caused high morality in C 3 grasses, which has large ecological and economic ramifications for this region. Furthermore, despite predictions based on the inverse texture hypothesis, we observed no interactive effects of soils with the drought treatment on cover or mortality. These results suggest that plant responses to droughts in drylands may differ from expectations and have large ecological effects if press- and pulse-droughts push species beyond physiological and mortality thresholds.

Oecologia

A case for using Plethodontid salamanders for monitoring biodiversity and ecosystem integrity of North American forests

Terrestrial salamanders of the family Plethodontidae have unique attributes that make them excellent indicators of biodiversity and ecosystem integrity in forested habitats. Their longevity, small territory size, site fidelity, sensitivity to natural and anthropogenic perturbations, tendency to occur in high densities, and low sampling costs mean that counts of plethodontid salamanders provide numerous advantages over counts of other North American forest organisms for indicating environmental change. Furthermore, they are tightly linked physiologically to microclimatic and successional processes that influence the distribution and abundance of numerous other hydrophilic but difficult-to-study forest-dwelling plants and animals. Ecosystem processes such as moisture cycling, food-web dynamics, and succession, with their related structural and microclimatic variability, all affect forest biodiversity and have been shown to affect salamander populations as well. We determined the variability associated with sampling for plethodontid salamanders by estimating the coefficient of variation (CV ) from available time-series data. The median coefficient of variation indicated that variation in counts of individuals among studies was much lower in plethodontids (27%) than in lepidoptera (93%), passerine birds (57%), small mammals (69%), or other amphibians (37–46%), which means plethodontid salamanders provide an important statistical advantage over other species for monitoring long-term forest health.

Conservation Biology

Effects of nitrogen deposition and empirical nitrogen critical loads for ecoregions of the United States

Human activity in the last century has led to a significant increase in nitrogen (N) emissions and atmospheric deposition. This N deposition has reached a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. One approach for quantifying the deposition of pollution that would be harmful to ecosystems is the determination of critical loads. A critical load is defined as the input of a pollutant below which no detrimental ecological effects occur over the long-term according to present knowledge. The objectives of this project were to synthesize current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and to estimate associated empirical N critical loads. The receptors considered included freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Ecosystem impacts included: (1) biogeochemical responses and (2) individual species, population, and community responses. Biogeochemical responses included increased N mineralization and nitrification (and N availability for plant and microbial uptake), increased gaseous N losses (ammonia volatilization, nitric and nitrous oxide from nitrification and denitrification), and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root : shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species. The range of critical loads for nutrient N reported for U.S. ecoregions, inland surface waters, and freshwater wetlands is 1-39 kg N.ha -1.yr -1, spanning the range of N deposition observed over most of the country. The empirical critical loads for N tend to increase in the following sequence for different life forms: diatoms, lichens and bryophytes, mycorrhizal fungi, herbaceous plants and shrubs, and trees. The critical load approach is an ecosystem assessment tool with great potential to simplify complex scientific information and communicate effectively with the policy community and the public. This synthesis represents the first comprehensive assessment of empirical critical loads of N for major ecoregions across the United States. ?? 2011 by the Ecological Society of America.

Ecological Applications

Growth and behavior of North American microbes on Phragmites australis leaves

Phragmites australis subsp. australis is a cosmopolitan wetland grass that is invasive in many regions of the world, including North America, where it co-occurs with the closely related Phragmites australis subsp. americanus. Because the difference in invasive behavior is unlikely to be related to physiological differences, we hypothesize that interactions with unique members of their microbiomes may significantly affect the behavior of each subspecies. Therefore, we systematically inoculated both plant lineages with a diverse array of 162 fungal and bacterial isolates to determine which could (1) differentiate between Phragmites hosts, (2) infect leaves at various stages of development, or (3) obtain plant-based carbon saprophytically. We found that many of the microbes isolated from Phragmites leaves behave as saprophytes. Only 1% (two taxa) were determined to be strong pathogens, 12% (20 taxa) were weakly pathogenic, and the remaining 87% were nonpathogenic. None of the isolates clearly discriminated between host plant lineages, and the Phragmites cuticle was shown to be a strong nonspecific barrier to infection. These results largely agree with the broad body of literature on leaf-associated phyllosphere microbes in Phragmites.

Microorganisms

Drivers of chaparral plant diversity

Chaparral diversity has marked spatial and temporal variation. Evolutionary diversity at the genetic, specific, and lineage level contribute to a very diverse flora. Ecological diversity is evident in life histories that comprise a range of physiological and morphological strategies for dealing with drought, and demographic patterns centered around different seedling recruitment strategies. Community or alpha diversity varies markedly through time. Mature chaparral ranges from monotypic stands of chamise ( Adenostoma fasciculatum ) to mixed chaparral often with up to a dozen shrub species. The understory contributes relatively little other than a few diminutive annuals and occasional herbaceous perennial resprouts. However, after fire, diversity increases dramatically and is often dominated by annuals that arise from a dormant seedbank with significant contribution of geophytes resprouting and flowering from dormant bulbs and corms. This flora has very diverse life histories, with some present only a year or two and then existing as a dormant seedbank or bulbs until the next fire. Others may persist much longer, often in gaps in the shrub canopy. Postfire dominance-diversity patterns fit a geometric model as most communities are dominated by a few species and the bulk of the flora comprise subordinates that occupy specific microhabitats. Postfire community assembly is a result of competitive interactions and environmental filtering effects. Beta diversity plays a role in community assembly for as heterogeneity of communities in the landscape increases, the potential species pool for a community increases. Gamma diversity is particularly high because species turnover across latitudinal and elevational gradients is high. The role of diversity in conferring community resilience is complex and a function of the life history of shrub dominants and the historical patterns of fires. Under some circumstances low diversity may be more resilient than high diversity, for example under high fire frequency monotypic stands of Adenostoma fasciculatum may resist change better than diverse stands that include obligate seeding shrubs sensitive to short interval fires. Postfire annuals also are sensitive to short interval fires as these disturbances enhance the invasion by more competitive non-native grasses. Expected increases in anthropogenic ignitions due to population growth are the biggest threat to biodiversity in chaparral.

Book chapter

Interactions across spatial scales among forest dieback, fire, and erosion in northern New Mexico landscapes

Ecosystem patterns and disturbance processes at one spatial scale often interact with processes at another scale, and the result of such cross-scale interactions can be nonlinear dynamics with thresholds. Examples of cross-scale pattern-process relationships and interactions among forest dieback, fire, and erosion are illustrated from northern New Mexico (USA) landscapes, where long-term studies have recently documented all of these disturbance processes. For example, environmental stress, operating on individual trees, can cause tree death that is amplified by insect mortality agents to propagate to patch and then landscape or even regional-scale forest dieback. Severe drought and unusual warmth in the southwestern USA since the late 1990s apparently exceeded species-specific physiological thresholds for multiple tree species, resulting in substantial vegetation mortality across millions of hectares of woodlands and forests in recent years. Predictions of forest dieback across spatial scales are constrained by uncertainties associated with: limited knowledge of species-specific physiological thresholds; individual and site-specific variation in these mortality thresholds; and positive feedback loops between rapidly-responding insect herbivore populations and their stressed plant hosts, sometimes resulting in nonlinear “pest” outbreak dynamics. Fire behavior also exhibits nonlinearities across spatial scales, illustrated by changes in historic fire regimes where patch-scale grazing disturbance led to regional-scale collapse of surface fire activity and subsequent recent increases in the scale of extreme fire events in New Mexico. Vegetation dieback interacts with fire activity by modifying fuel amounts and configurations at multiple spatial scales. Runoff and erosion processes are also subject to scale-dependent threshold behaviors, exemplified by ecohydrological work in semiarid New Mexico watersheds showing how declines in ground surface cover lead to non-linear increases in bare patch connectivity and thereby accelerated runoff and erosion at hillslope and watershed scales. Vegetation dieback, grazing, and fire can change land surface properties and cross-scale hydrologic connectivities, directly altering ecohydrological patterns of runoff and erosion. The interactions among disturbance processes across spatial scales can be key drivers in ecosystem dynamics, as illustrated by these studies of recent landscape changes in northern New Mexico. To better anticipate and mitigate accelerating human impacts to the planetary ecosystem at all spatial scales, improvements are needed in our conceptual and quantitative understanding of cross-scale interactions among disturbance processes.

New Mexico