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Societal challenges in understanding and responding to regime shifts in forest landscapes

2 excerpts: "The degradation of seminatural landscapes at regional scales, whereby essential functional capabilities and biotic elements are permanently lost as a result of altered disturbance regimes, is a widespread phenomenon." and "Salvage logging of burned or windthrown forests not only eliminates critical structural legacies from predisturbance stands but can disrupt natural regenerative processes, as noted below (10, 11)."

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Genome diversity in wild grasses under environmental stress

Patterns of diversity distribution in the Isa defense locus in wild- barley populations suggest adaptive selection at this locus. The extent to which environmental selection may act at additional nuclear-encoded defense loci and within the whole chloroplast genome has now been examined by analyses in two grass species. Analysis of genetic diversity in wild barley (Hordeum spontaneum) defense genes revealed much greater variation in biotic stress-related genes than abiotic stress-related genes. Genetic diversity at the Isa defense locus in wild populations of weeping ricegrass [Microlaena stipoides (Labill.) R. Br.], a very distant wild-rice relative, was more diverse in samples from relatively hotter and drier environments, a phenomenon that reflects observations in wild barley populations. Whole-chloroplast genome sequences of bulked weeping ricegrass individuals sourced from contrasting environments showed higher levels of diversity in the drier environment in both coding and noncoding portions of the genome. Increased genetic diversity may be important in allowing plant populations to adapt to greater environmental variation in warmer and drier climatic conditions. adaptive variation | genomics | molecular evolution | disease resistance | abiotic stress resistance

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Satellite-based global-ocean mass balance estimates of interannual variability and emerging trends in continental freshwater discharge

Freshwater discharge from the continents is a key component of Earth’s water cycle that sustains human life and ecosystem health. Surprisingly, owing to a number of socioeconomic and political obstacles, a comprehensive global river discharge observing system does not yet exist. Here we use 13 years (1994–2006) of satellite precipitation, evaporation, and sea level data in an ocean mass balance to estimate freshwater discharge into the global ocean. Results indicate that global freshwater discharge averaged 36,055 km3∕y for the study period while exhibiting significant interannual variability driven primarily by El Niño Southern Oscillation cycles. The method described here can ultimately be used to estimate long-term global discharge trends as the records of sea level rise and ocean temperature lengthen. For the relatively short 13-year period studied here, global discharge increased by 540 km3 ∕y2 , which was largely attributed to an increase of global- ocean evaporation (768 km3 ∕y2 ). Sustained growth of these flux rates into long-term trends would provide evidence for increasing intensity of the hydrologic cycle. climate ∣ global water cycle ∣ hydrology ∣ remote sensing ∣ observations

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Temperature increase of 21st century mitigation scenarios

Estimates of 21st Century global-mean surface temperature in- crease have generally been based on scenarios that do not include climate policies. Newly developed multigas mitigation scenarios, based on a wide range of modeling approaches and socioeconomic assumptions, now allow the assessment of possible impacts of climate policies on projected warming ranges. This article assesses the atmospheric CO2 concentrations, radiative forcing, and tem- perature increase for these new scenarios using two reduced- complexity climate models. These scenarios result in temperature increase of 0.5–4.4°C over 1990 levels or 0.3–3.4°C less than the no-policy cases. The range results from differences in the assumed stringency of climate policy and uncertainty in our understanding of the climate system. Notably, an average minimum warming of 􏰂1.4°C (with a full range of 0.5–2.8°C) remains for even the most stringent stabilization scenarios analyzed here. This value is sub- stantially above previously estimated committed warming based on climate system inertia alone. The results show that, although ambitious mitigation efforts can significantly reduce global warming, adaptation measures will be needed in addition to mitigation to reduce the impact of the residual warming. climate 􏰀 climate policy 􏰀 stabilization 􏰀 integrated assessment 􏰀 scenario

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Impacts of climate change on the world’s most exceptional ecoregions

The current rate of warming due to increases in greenhouse gas (GHG) emissions is very likely unprecedented over the last 10,000 y. Although the majority of countries have adopted the view that global warming must be limited to <2 °C, current GHG emission rates and nonagreement at Copenhagen in December 2009 increase the likelihood of this limit being exceeded by 2100. Extensive evi- dence has linked major changes in biological systems to 20th century warming. The “Global 200” comprises 238 ecoregions of exceptional biodiversity [Olson DM, Dinerstein E (2002) Ann Mo Bot Gard 89:199–224]. We assess the likelihood that, by 2070, these iconic ecoregions will regularly experience monthly climatic conditions that were extreme in 1961–1990. Using >600 realizations from climate model ensembles, we show that up to 86% of terres- trial and 83% of freshwater ecoregions will be exposed to average monthly temperature patterns >2 SDs (2σ) of the 1961–1990 base- line, including 82% of critically endangered ecoregions. The entire range of 89 ecoregions will experience extreme monthly temper- atures with a local warming of <2 °C. Tropical and subtropical ecor- egions, and mangroves, face extreme conditions earliest, some with <1 °C warming. In contrast, few ecoregions within Boreal Forests and Tundra biomes will experience such extremes this cen- tury. On average, precipitation regimes do not exceed 2σ of the baseline period, although considerable variability exists across the climate realizations. Further, the strength of the correlation between seasonal temperature and precipitation changes over nu- merous ecoregions. These results suggest many Global 200 ecore- gions may be under substantial climatic stress by 2100. climate impacts | climate model ensemble | conservation extreme

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Protected areas facilitate species’ range expansions

The benefits of protected areas (PAs) for biodiversity have been questioned in the context of climate change because PAs are static, whereas the distributions of species are dynamic. Current PAs may, however, continue to be important if they provide suitable locations for species to colonize at their leading-edge range boundaries, thereby enabling spread into new regions. Here, we present an empirical assessment of the role of PAs as targets for coloniza- tion during recent range expansions. Records from intensive sur- veys revealed that seven bird and butterfly species have colonized PAs 4.2 (median) times more frequently than expected from the availability of PAs in the landscapes colonized. Records of an additional 256 invertebrate species with less-intensive surveys supported these findings and showed that 98% of species are disproportionately associated with PAs in newly colonized parts of their ranges. Although colonizing species favor PAs in general, species vary greatly in their reliance on PAs, reflecting differences in the dependence of individual species on particular habitats and other conditions that are available only in PAs. These findings highlight the importance of current PAs for facilitating range expansions and show that a small subset of the landscape receives a high proportion of colonizations by range-expanding species. conservation | climate change adaptation | nature reserves

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Soil warming, carbon–nitrogen interactions, and forest carbon budgets

Soil warming has the potential to alter both soil and plant processes that affect carbon storage in forest ecosystems. We have quantified these effects in a large, long-term (7-y) soil-warming study in a deciduous forest in New England. Soil warming has resulted in carbon losses from the soil and stimulated carbon gains in the woody tissue of trees. The warming-enhanced decay of soil organic matter also released enough additional inorganic nitrogen into the soil solution to support the observed increases in plant carbon storage. Although soil warming has resulted in a cumulative net loss of carbon from a New England forest relative to a control area over the 7-y study, the annual net losses generally decreased over time as plant carbon storage increased. In the seventh year, warming-induced soil carbon losses were almost totally compensated for by plant carbon gains in response to warming. We attribute the plant gains primarily to warming- induced increases in nitrogen availability. This study underscores the importance of incorporating carbon–nitrogen interactions in atmosphere–ocean–land earth system models to accurately simulate land feedbacks to the climate system. climate system feedbacks | ecological stoichiometry | forest carbon budget | forest nitrogen budget | global climate change

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Latitudinal variation in lifespan within species is explained by the metabolic theory of ecology

Many ectotherms exhibit striking latitudinal gradients in lifespan. However, it is unclear whether lifespan gradients in distantly related taxa share a common mechanistic explanation. We com- piled data on geographic variation in lifespan in ectotherms from around the globe to determine how much of this intraspecific variation in lifespan may be explained by temperature using the simple predictions of the metabolic theory of ecology. We found that the metabolic theory accurately predicts how lifespan varies with temperature within species in a wide range of ectotherms in both controlled laboratory experiments and free-living populations. After removing the effect of temperature, only a small fraction of species showed significant trends with latitude. There was, however, considerable residual intraspecific variation indi- cating that other, more local factors are likely to be important in determining lifespan within species. These findings suggest that, given predicted increases in global temperature, lifespan of ectotherms may be substantially shortened in the future. ectotherms 􏰀 intraspecific 􏰀 longevity 􏰀 MTE

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Global warming benefits the small in aquatic ecosystems

Understanding the ecological impacts of climate change is a crucial challenge of the twenty-first century. There is a clear lack of general rules regarding the impacts of global warming on biota. Here, we present a metaanalysis of the effect of climate change on body size of ectothermic aquatic organisms (bacteria, phyto- and zooplankton, and fish) from the community to the individual level. Using long-term surveys, experimental data and published results, we show a significant increase in the proportion of small-sized species and young age classes and a decrease in size-at-age. These results are in accordance with the ecological rules dealing with the temperature–size relationships (i.e., Bergmann’s rule, James’ rule and Temperature–Size Rule). Our study provides evidence that reduced body size is the third universal ecological response to global warming in aquatic systems besides the shift of species ranges toward higher altitudes and latitudes and the seasonal shifts in life cycle events. biological scale 􏰀 body size 􏰀 climate change 􏰀 ectotherms 􏰀 metaanalysis

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Nonlinear temperature effects indicate severe damages to U.S. crop yields under climate change

The United States produces 41% of the world’s corn and 38% of the world’s soybeans. These crops comprise two of the four largest sources of caloric energy produced and are thus critical for world food supply. We pair a panel of county-level yields for these two crops, plus cotton (a warmer-weather crop), with a new fine-scale weather dataset that incorporates the whole distribution of tem- peratures within each day and across all days in the growing season. We find that yields increase with temperature up to 29° C for corn, 30° C for soybeans, and 32° C for cotton but that tem- peratures above these thresholds are very harmful. The slope of the decline above the optimum is significantly steeper than the incline below it. The same nonlinear and asymmetric relationship is found when we isolate either time-series or cross-sectional variations in temperatures and yields. This suggests limited his- torical adaptation of seed varieties or management practices to warmer temperatures because the cross-section includes farmers’ adaptations to warmer climates and the time-series does not. Holding current growing regions fixed, area-weighted average yields are predicted to decrease by 30 – 46% before the end of the century under the slowest (B1) warming scenario and decrease by 63–82% under the most rapid warming scenario (A1FI) under the Hadley III model. agriculture 􏰀 panel analysis 􏰀 time series 􏰀 cross section 􏰀 farmer adaptation

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Temperature sensitivity of drought-induced tree mortality portends increased regional die-off under global change-type drought

Large-scale biogeographical shifts in vegetation are predicted in response to the altered precipitation and temperature regimes associated with global climate change. Vegetation shifts have profound ecological impacts and are an important climate-ecosystem feedback through their alteration of carbon, water, and energy exchanges of the land surface. Of particular concern is the potential for warmer temperatures to compound the effects of increasingly severe droughts by triggering widespread vegetation shifts via woody plant mortality. The sensitivity of tree mortality to temperature is dependent on which of 2 non-mutually-exclusive mechanisms predominates—temperature-sensitive carbon starvation in response to a period of protracted water stress or temperature-insensitive sudden hydraulic failure under extreme water stress (cavitation). Here we show that experimentally induced warmer temperatures (􏰂4 °C) shortened the time to drought- induced mortality in Pinus edulis (pin ̃ on shortened pine) trees by nearly a third, with temperature-dependent differences in cumu- lative respiration costs implicating carbon starvation as the primary mechanism of mortality. Extrapolating this temperature effect to the historic frequency of water deficit in the southwestern United States predicts a 5-fold increase in the frequency of regional-scale tree die-off events for this species due to temperature alone. Projected increases in drought frequency due to changes in pre- cipitation and increases in stress from biotic agents (e.g., bark beetles) would further exacerbate mortality. Our results demon- strate the mechanism by which warmer temperatures have exac- erbated recent regional die-off events and background mortality rates. Because of pervasive projected increases in temperature, our results portend widespread increases in the extent and frequency of vegetation die-off. biosphere–atmosphere feedbacks 􏰀 drought impacts 􏰀 global-change ecology 􏰀 Pinus edulis 􏰀 carbon starvation

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Maximum leaf conductance driven by CO2 effects on stomatal size and density over geologic time

Stomatal pores are microscopic structures on the epidermis of leaves formed by 2 specialized guard cells that control the exchange of water vapor and CO2 between plants and the atmosphere. Stomatal size (S) and density (D) determine maximum leaf diffusive (stomatal) conductance of CO2 (gcmax) to sites of assimi- lation. Although large variations in D observed in the fossil record have been correlated with atmospheric CO2, the crucial significance of similarly large variations in S has been overlooked. Here, we use physical diffusion theory to explain why large changes in S nec- essarily accompanied the changes in D and atmospheric CO2 over the last 400 million years. In particular, we show that high densities of small stomata are the only way to attain the highest gcmax values required to counter CO2‘‘starvation’’ at low atmospheric CO2 concentrations. This explains cycles of increasing D and decreasing S evident in the fossil history of stomata under the CO2 impover- ished atmospheres of the Permo-Carboniferous and Cenozoic gla- ciations. The pattern was reversed under rising atmospheric CO2 regimes. Selection for small S was crucial for attaining high gcmax under falling atmospheric CO2 and, therefore, may represent a mechanism linking CO2 and the increasing gas-exchange capacity of land plants over geologic time. Phanerozoic 􏰀 photosynthesis 􏰀 plant evolution 􏰀 transpiration 􏰀 xylem

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Irreversible climate change due to carbon dioxide emissions

The severity of damaging human-induced climate change depends not only on the magnitude of the change but also on the potential for irreversibility. This paper shows that the climate change that takes place due to increases in carbon dioxide concentration is largely irreversible for 1,000 years after emissions stop. Following cessation of emissions, removal of atmospheric carbon dioxide decreases radiative forcing, but is largely compensated by slower loss of heat to the ocean, so that atmospheric temperatures do not drop significantly for at least 1,000 years. Among illustrative irreversible impacts that should be expected if atmospheric carbon dioxide concentrations increase from current levels near 385 parts per million by volume (ppmv) to a peak of 450 – 600 ppmv over the coming century are irreversible dry-season rainfall reductions in several regions comparable to those of the ‘‘dust bowl’’ era and inexorable sea level rise. Thermal expansion of the warming ocean provides a conservative lower limit to irreversible global average sea level rise of at least 0.4–1.0 m if 21st century CO2 concentrations exceed 600 ppmv and 0.6 –1.9 m for peak CO2 concentrations exceeding 1,000 ppmv. Additional contributions from glaciers and ice sheet contributions to future sea level rise are uncertain but may equal or exceed several meters over the next millennium or longer. dangerous interference 􏰀 precipitation 􏰀 sea level rise 􏰀 warming

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Overcoming systemic roadblocks to sustainability: The evolutionary redesign of worldviews, institutions, and technologies

A high and sustainable quality of life is a central goal for humanity. Our current socio-ecological regime and its set of interconnected worldviews, institutions, and technologies all support the goal of unlimited growth of material production and consumption as a proxy for quality of life. However, abundant evidence shows that, beyond a certain threshold, further material growth no longer significantly contributes to improvement in quality of life. Not only does further material growth not meet humanity’s central goal, there is mounting evidence that it creates significant roadblocks to sustainability through increasing resource constraints (i.e., peak oil, water limitations) and sink constraints (i.e., climate disruption). Overcoming these roadblocks and creating a sustainable and de- sirable future will require an integrated, systems level redesign of our socio-ecological regime focused explicitly and directly on the goal of sustainable quality of life rather than the proxy of unlimited material growth. This transition, like all cultural transitions, will occur through an evolutionary process, but one that we, to a certain extent, can control and direct. We suggest an integrated set of worldviews, institutions, and technologies to stimulate and seed this evolutionary redesign of the current socio-ecological regime to achieve global sustainability. cultural adaptation 􏰀 ecology 􏰀 societal decline

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The physical basis for increases in precipitation extremes in simulations of 21st-century climate change

Global warming is expected to lead to a large increase in atmospheric water vapor content and to changes in the hydrological cycle, which include an intensification of precipitation extremes. The intensity of precipitation extremes is widely held to increase proportionately to the increase in atmospheric water vapor content. Here, we show that this is not the case in 21st-century climate change scenarios simulated with climate models. In the tropics, precipitation extremes are not simulated reliably and do not change consistently among climate models; in the extratropics, they consistently increase more slowly than atmospheric water vapor content. We give a physical basis for how precipitation extremes change with climate and show that their changes depend on changes in the moist-adiabatic temperature lapse rate, in the upward velocity, and in the temperature when precipitation ex- tremes occur. For the tropics, the theory suggests that improving the simulation of upward velocities in climate models is essential for improving predictions of precipitation extremes; for the extra- tropics, agreement with theory and the consistency among climate models increase confidence in the robustness of predictions of precipitation extremes under climate change. global warming 􏰀 hydrological cycle 􏰀 rainfall 􏰀 extreme events

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Synchronous extinction of North America’s Pleistocene mammals

The late Pleistocene witnessed the extinction of 35 genera of North American mammals. The last appearance dates of 16 of these genera securely fall between 12,000 and 10,000 radiocarbon years ago (􏰂13,800–11,400 calendar years B.P.), although whether the absence of fossil occurrences for the remaining 19 genera from this time interval is the result of sampling error or temporally staggered extinctions is unclear. Analysis of the chronology of extinctions suggests that sampling error can explain the absence of terminal Pleistocene last appearance dates for the remaining 19 genera. The extinction chronology of North American Pleistocene mammals therefore can be characterized as a synchronous event that took place 12,000–10,000 radiocarbon years B.P. Results favor an ex- tinction mechanism that is capable of wiping out up to 35 genera across a continent in a geologic instant. climate change 􏰀 extraterrestrial impact 􏰀 overkill 􏰀 Quaternary extinctions 􏰀 radiocarbon dates

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Multidimensional evaluation of managed relocation

Managed relocation (MR) has rapidly emerged as a potential intervention strategy in the toolbox of biodiversity management under climate change. Previous authors have suggested that MR (also referred to as assisted colonization, assisted migration, or assisted translocation) could be a last-alternative option after interrogating a linear decision tree. We argue that numerous interacting and value-laden considerations demand a more inclu- sive strategy for evaluating MR. The pace of modern climate change demands decision making with imperfect information, and tools that elucidate this uncertainty and integrate scientific information and social values are urgently needed. We present a heuristic tool that incorporates both ecological and social criteria in a multidimensional decision-making framework. For visualization purposes, we collapse these criteria into 4 classes that can be depicted in graphical 2-D space. This framework offers a pragmatic approach for summarizing key dimensions of MR: capturing un- certainty in the evaluation criteria, creating transparency in the evaluation process, and recognizing the inherent tradeoffs that different stakeholders bring to evaluation of MR and its alternatives. assisted migration 􏰀 climate change 􏰀 conservation biology 􏰀 conservation strategy 􏰀 sustainability science

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Re-evaluation of forest biomass carbon stocks and lessons from the world’s most carbon-dense forests

From analysis of published global site biomass data (n = 136) from primary forests, we discovered (i) the world’s highest known total biomass carbon density (living plus dead) of 1,867 tonnes carbon per ha (average value from 13 sites) occurs in Australian temperate moist Eucalyptus regnans forests, and (ii) average values of the global site biomass data were higher for sampled temperate moist forests (n 􏰀 44) than for sampled tropical (n 􏰀 36) and boreal (n 􏰀 52) forests (n is number of sites per forest biome). Spatially averaged Intergovern- mental Panel on Climate Change biome default values are lower than our average site values for temperate moist forests, because the temperate biome contains a diversity of forest ecosystem types that support a range of mature carbon stocks or have a long land-use history with reduced carbon stocks. We describe a framework for identifying forests important for carbon storage based on the factors that account for high biomass carbon densities, including (i) relatively cool temperatures and moderately high precipitation producing rates of fast growth but slow decomposition, and (ii) older forests that are often multiaged and multilayered and have experienced minimal human disturbance. Our results are relevant to negotiations under the United Nations Framework Convention on Climate Change re- garding forest conservation, management, and restoration. Conserv- ing forests with large stocks of biomass from deforestation and degradation avoids significant carbon emissions to the atmosphere, irrespective of the source country, and should be among allowable mitigation activities. Similarly, management that allows restoration of a forest’s carbon sequestration potential also should be recognized. Eucalyptus regnans 􏰂 climate mitigation 􏰂 primary forest 􏰂 deforestation and degradation 􏰂 temperate moist forest biome

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The potential for behavioral thermoregulation to buffer ‘‘cold-blooded’’ animals against climate warming

Increasing concern about the impacts of global warming on biodi- versity has stimulated extensive discussion, but methods to trans- late broad-scale shifts in climate into direct impacts on living animals remain simplistic. A key missing element from models of climatic change impacts on animals is the buffering influence of behavioral thermoregulation. Here, we show how behavioral and mass/energy balance models can be combined with spatial data on climate, topography, and vegetation to predict impacts of in- creased air temperature on thermoregulating ectotherms such as reptiles and insects (a large portion of global biodiversity). We show that for most ‘‘cold-blooded’’ terrestrial animals, the primary thermal challenge is not to attain high body temperatures (al- though this is important in temperate environments) but to stay cool (particularly in tropical and desert areas, where ectotherm biodiversity is greatest). The impact of climate warming on ther- moregulating ectotherms will depend critically on how changes in vegetation cover alter the availability of shade as well as the animals’ capacities to alter their seasonal timing of activity and reproduction. Warmer environments also may increase mainte- nance energy costs while simultaneously constraining activity time, putting pressure on mass and energy budgets. Energy- and mass-balance models provide a general method to integrate the complexity of these direct interactions between organisms and climate into spatial predictions of the impact of climate change on biodiversity. This methodology allows quantitative organism- and habitat-specific assessments of climate change impacts. Australia 􏰚 biophysical model 􏰚 climate change 􏰚 terrestrial ectotherm 􏰚 GIS

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The future of ice sheets and sea ice: Between reversible retreat and unstoppable loss

We discuss the existence of cryospheric “tipping points” in the Earth’s climate system. Such critical thresholds have been sug- gested to exist for the disappearance of Arctic sea ice and the retreat of ice sheets: Once these ice masses have shrunk below an anticipated critical extent, the ice–albedo feedback might lead to the irreversible and unstoppable loss of the remaining ice. We here give an overview of our current understanding of such thresh- old behavior. By using conceptual arguments, we review the recent findings that such a tipping point probably does not exist for the loss of Arctic summer sea ice. Hence, in a cooler climate, sea ice could recover rapidly from the loss it has experienced in recent years. In addition, we discuss why this recent rapid retreat of Arc- tic summer sea ice might largely be a consequence of a slow shift in ice-thickness distribution, which will lead to strongly increased year-to-year variability of the Arctic summer sea-ice extent. This variability will render seasonal forecasts of the Arctic summer sea- ice extent increasingly difficult. We also discuss why, in contrast to Arctic summer sea ice, a tipping point is more likely to exist for the loss of the Greenland ice sheet and the West Antarctic ice sheet. Greenland | West Antarctic | climate change | tipping point | Arctic

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