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A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests

Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. Increases in the frequency, duration, and/or severity of drought and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography of forests in many regions. Of particular concern are potential increases in tree mortality associated with climate-induced physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite this risk, existing projections of tree mortality are based on models that lack functionally realistic mortality mechanisms, and there has been no attempt to track observations of climate-driven tree mortality globally. Here we present the first global assessment of recent tree mortality attributed to drought and heat stress. Although episodic mortality occurs in the absence of climate change, studies compiled here suggest that at least some of the world's forested ecosystems already may be responding to climate change and raise concern that forests may become increasingly vulnerable to higher background tree mortality rates and die-off in response to future warming and drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem services, including the loss of sequestered forest carbon and associated atmospheric feedbacks. Our review also identifies key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the potential for amplified tree mortality due to drought and heat in forests worldwide.

Forest Ecology and Management

Benthic community metrics track hydrologically stressed mangrove systems

Mangrove restoration efforts have increased in order to help combat their decline globally. While restoration efforts often focus on planting seedlings, underlying chronic issues, including disrupted hydrological regimes, can hinder restoration success. While improving hydrology may be more cost-effective and have higher success rates than planting seedlings alone, hydrological restoration success in this form is poorly understood. Restoration assessments can employ a functional equivalency approach, comparing restoration areas over time with natural, reference forests in order to quantify the relative effectiveness of different restoration approaches. Here, we employ the use of baseline community ecology metrics along with stable isotopes to track changes in the community and trophic structure and enable time estimates for establishing mangrove functional equivalency. We examined a mangrove system impacted by road construction and recently targeted for hydrological restoration within the Rookery Bay National Estuarine Research Reserve, Florida, USA. Samples were collected along a gradient of degradation, from a heavily degraded zone, with mostly dead trees, to a transition zone, with a high number of saplings, to a full canopy zone, with mature trees, and into a reference zone with dense, mature mangrove trees. The transition, full canopy, and reference zones were dominated by annelids, gastropods, isopods, and fiddler crabs. Diversity was lower in the dead zone; these taxa were enriched in 13 C relative to those found in all the other zones, indicating a shift in the dominant carbon source from mangrove detritus (reference zone) to algae (dead zone). Community-wide isotope niche metrics also distinguished zones, likely reflecting dominant primary food resources (baseline organic matter) present. Our results suggest that stable isotope niche metrics provide a useful tool for tracking mangrove degradation gradients. These baseline data provide critical information on the ecosystem functioning in mangrove habitats following hydrological restoration.

Florida

Spectral mixture analyses of hyperspectral data acquired using a tethered balloon

Tethered balloon remote sensing platforms can be used to study radiometric issues in terrestrial ecosystems by effectively bridging the spatial gap between measurements made on the ground and those acquired via airplane or satellite. In this study, the Short Wave Aerostat-Mounted Imager (SWAMI) tethered balloon-mounted platform was utilized to evaluate linear and nonlinear spectral mixture analysis (SMA) for a grassland-conifer forest ecotone during the summer of 2003. Hyperspectral measurement of a 74-m diameter ground instantaneous field of view (GIFOV) attained by the SWAMI was studied. Hyperspectral spectra of four common endmembers, bare soil, grass, tree, and shadow, were collected in situ, and images captured via video camera were interpreted into accurate areal ground cover fractions for evaluating the mixture models. The comparison between the SWAMI spectrum and the spectrum derived by combining in situ spectral data with video-derived areal fractions indicated that nonlinear effects occurred in the near infrared (NIR) region, while nonlinear influences were minimal in the visible region. The evaluation of hyperspectral and multispectral mixture models indicated that nonlinear mixture model-derived areal fractions were sensitive to the model input data, while the linear mixture model performed more stably. Areal fractions of bare soil were overestimated in all models due to the increased radiance of bare soil resulting from side scattering of NIR radiation by adjacent grass and trees. Unmixing errors occurred mainly due to multiple scattering as well as close endmember spectral correlation. In addition, though an apparent endmember assemblage could be derived using linear approaches to yield low residual error, the tree and shade endmember fractions calculated using this technique were erroneous and therefore separate treatment of endmembers subject to high amounts of multiple scattering (i.e. shadows and trees) must be done with caution. Including the short wave infrared (SWIR) region in the hyperspectral and multispectral endmember data significantly reduced the Pearson correlation coefficient values among endmember spectra. Therefore, combination of visible, NIR, and SWIR information is likely to further improve the utility of SMA in understanding ecosystem structure and function and may help narrow uncertainties when utilizing remotely sensed data to extrapolate trace glas flux measurements from the canopy scale to the landscape scale.

Remote Sensing of Environment

Passive restoration following ungulate removal in a highly disturbed tropical wet forest devoid of native seed dispersers

Overabundant ungulate populations can alter forests. Concurrently, global declines of seed dispersers may threaten native forest structure and function. On an island largely devoid of native vertebrate seed dispersers, we monitored forest succession for 7 years following ungulate exclusion from a 5-ha area and adjacent plots with ungulates still present. We observed succession from open scrub to forest and understory cover by non-native plants declined. Two trees, native Hibiscus tiliaceus and non-native Leucaena leucocephala , accounted for most forest regeneration, with the latter dominant. Neither species is dependent on animal dispersers nor was there strong evidence that plants dependent on dispersers migrated into the 5-ha study area. Passive restoration following ungulate removal may facilitate restoration, but did not show promise for fully restoring native forest on Guam. Restoration of native forest plants in bird depopulated areas will likely require active outplanting of native seedlings, control of factors resulting in bird loss, and reintroduction of seed dispersers.

Guam

Nutrient controls on biocomplexity of mangrove ecosystems

Mangrove forests are important coastal ecosystems that provide a variety of ecological and societal services. These intertidal, tree-dominated communities along tropical coastlines are often described as “simple systems,” compared to other tropical forests with larger numbers of plant species and multiple understory strata; however, mangrove ecosystems have complex trophic structures, and organisms exhibit unique physiological, morphological, and behavioral adaptations to environmental conditions characteristic of the land-sea interface. Biogeochemical functioning of mangrove forests is also controlled by interactions among the microbial, plant, and animal communities and feedback linkages mediated by hydrology and other forcing functions. Scientists with the U.S. Geological Survey (USGS) at the National Wetlands Research Center are working to understand more fully the impact of nutrient variability on these delicate and important ecosystems.

Fact Sheet

Sagebrush recovery patterns after fuel treatments mediated by disturbance type and plant functional group interactions

Fire and fuel management is a high priority in North American sagebrush ecosystems where the expansion of piñon and juniper trees and the invasion of nonnative annual grasses are altering fire regimes and resulting in loss of sagebrush species and habitat. We evaluated 10-yr effects of woody fuel treatments on sagebrush recruitment and plant functional group interactions using Sagebrush Steppe Treatment Evaluation Project data. We used mixed-effects ANOVAs to examine treatment effects on sagebrush density and cover and perennial and annual grass cover in expansion woodlands (prescribed fire and cut-and-leave) and annual grass invasion areas (prescribed fire, mowing, tebuthiuron herbicide application). We used piecewise structural equation models to evaluate interactions among sagebrush seedling density, juvenile and adult density, and cover and perennial and annual grass cover. Fuel treatments were equated to pulse or press disturbances varying in resource release and subsequent intra- and interspecific interactions. Prescribed fire, a high magnitude pulse disturbance with more severe effects in warm and dry sites, reduced sagebrush cover and decoupled associations among sagebrush seedlings, juvenile and adult density, and cover indicating changed population structure. Cutting and leaving trees, a low magnitude pulse disturbance in cooler and moister woodlands, increased sagebrush density and cover and generally had lesser effects on sagebrush intraspecific associations. Mowing, a moderate magnitude pulse disturbance, and tebuthiuron herbicide application, a multiyear press disturbance, reduced sagebrush cover and disrupted intraspecific relationships. Competitive release increased cover of perennial grass in all treatments but tebuthiuron. Annual grass increased in all treatments, especially prescribed fire and tebuthiuron. Annual and perennial grass interactions with sagebrush were generally rare, but in woodland treatments perennial grass suppressed annual grass through year 6. Treatments in cooler and moister woodland sites had more positive effects on sagebrush recruitment and perennial grass cover, less negative effects on sagebrush intraspecific interactions, and smaller increases in annual grass cover indicating potential increases in resilience to fire. In warmer and drier invasion sites, reductions in woody fuels resulted in lack of sagebrush recruitment, disruption of sagebrush intraspecific interactions, and progressive increases in annual grass indicating reduced resilience to fire and resistance to invaders.

Ecosphere

Hurricane disturbance and recovery of energy balance, CO 2 fluxes and canopy structure in a mangrove forest of the Florida Everglades

Eddy covariance (EC) estimates of carbon dioxide (CO 2 ) fluxes and energy balance are examined to investigate the functional responses of a mature mangrove forest to a disturbance generated by Hurricane Wilma on October 24, 2005 in the Florida Everglades. At the EC site, high winds from the hurricane caused nearly 100% defoliation in the upper canopy and widespread tree mortality. Soil temperatures down to -50 cm increased, and air temperature lapse rates within the forest canopy switched from statically stable to statically unstable conditions following the disturbance. Unstable conditions allowed more efficient transport of water vapor and CO 2 from the surface up to the upper canopy layer. Significant increases in latent heat fluxes (LE) and nighttime net ecosystem exchange (NEE) were also observed and sensible heat fluxes (H) as a proportion of net radiation decreased significantly in response to the disturbance. Many of these impacts persisted through much of the study period through 2009. However, local albedo and MODIS (Moderate Resolution Imaging Spectro-radiometer) data (the Enhanced Vegetation Index) indicated a substantial proportion of active leaf area recovered before the EC measurements began 1 year after the storm. Observed changes in the vertical distribution and the degree of clumping in newly emerged leaves may have affected the energy balance. Direct comparisons of daytime NEE values from before the storm and after our measurements resumed did not show substantial or consistent differences that could be attributed to the disturbance. Regression analyses on seasonal time scales were required to differentiate the storm's impact on monthly average daytime NEE from the changes caused by interannual variability in other environmental drivers. The effects of the storm were apparent on annual time scales, and CO 2 uptake remained approximately 250 g C m -2 yr -1 lower in 2009 compared to the average annual values measured in 2004–2005. Dry season CO 2 uptake was relatively more affected by the disturbance than wet season values. Complex leaf regeneration dynamics on damaged trees during ecosystem recovery are hypothesized to lead to the variable dry versus wet season impacts on daytime NEE. In contrast, nighttime CO 2 release (i.e., nighttime respiration) was consistently and significantly greater, possibly as a result of the enhanced decomposition of litter and coarse woody debris generated by the storm, and this effect was most apparent in the wet seasons compared to the dry seasons. The largest pre- and post-storm differences in NEE coincided roughly with the delayed peak in cumulative mortality of stems in 2007–2008. Across the hurricane-impacted region, cumulative tree mortality rates were also closely correlated with declines in peat surface elevation. Mangrove forest–atmosphere interactions are interpreted with respect to the damage and recovery of stand dynamics and soil accretion processes following the hurricane.

Florida

Mummy Lake: An unroofed ceremonial structure within a large-scale ritual landscape

The structure at Mesa Verde National Park known historically as Mummy Lake and more recently as Far View Reservoir is not part of a water collection, impoundment, or redistribution system. We offer an alternative explanation for the function of Mummy Lake. We suggest that it is an unroofed ceremonial structure, and that it serves as an essential component of a Chacoan ritual landscape. A wide constructed avenue articulates Mummy Lake with Far View House and Pipe Shrine House. The avenue continues southward for approximately 6 km where it apparently divides connecting with Spruce Tree House and Sun Temple/Cliff Palace. The avenue has previously been interpreted as an irrigation ditch fed by water impounded at Mummy Lake; however, it conforms in every respect to alignments described as Chacoan roads. Tree-ring dates indicate that the construction of Spruce Tree House and Cliff Palace began about A.D. 1225, roughly coincident with the abandonment of the Far View community. This pattern of periodically relocating the focus of an Anasazi community by retiring existing ritual structures and linking them to newly constructed facilities by means of broad avenues was first documented by Fowler and Stein (1992) in Manuelito Canyon, New Mexico. Periods of intense drought appear to have contributed to the relocation of prehistoric Native Americans from the Far View group to Cliff Palace/Spruce Tree House in the mid-13th century and eventually to the abandonment of all Anasazi communities in southwestern Colorado in the late-13th century.

Journal of Archaeological Science

Biocomplexity in mangrove ecosystems

Mangroves are an ecological assemblage of trees and shrubs adapted to grow in intertidal environments along tropical coasts. Despite repeated demonstration of their economic and societal value, more than 50% of the world's mangroves have been destroyed, 35% in the past two decades to aquaculture and coastal development, altered hydrology, sea-level rise, and nutrient overenrichment. Variations in the structure and function of mangrove ecosystems have generally been described solely on the basis of a hierarchical classification of the physical characteristics of the intertidal environment, including climate, geomorphology, topography, and hydrology. Here, we use the concept of emergent properties at multiple levels within a hierarchical framework to review how the interplay between specialized adaptations and extreme trait plasticity that characterizes mangroves and intertidal environments gives rise to the biocomplexity that distinguishes mangrove ecosystems. The traits that allow mangroves to tolerate variable salinity, flooding, and nutrient availability influence ecosystem processes and ultimately the services they provide. We conclude that an integrated research strategy using emergent properties in empirical and theoretical studies provides a holistic approach for understanding and managing mangrove ecosystems.

Annual Review of Marine Science

Standing crop and aboveground biomass partitioning of a dwarf mangrove forest in Taylor River Slough, Florida

The structure and standing crop biomass of a dwarf mangrove forest, located in the salinity transition zone ofTaylor River Slough in the Everglades National Park, were studied. Although the four mangrove species reported for Florida occurred at the study site, dwarf Rhizophora mangle trees dominated the forest. The structural characteristics of the mangrove forest were relatively simple: tree height varied from 0.9 to 1.2 meters, and tree density ranged from 7062 to 23 778 stems haa??1. An allometric relationship was developed to estimate leaf, branch, prop root, and total aboveground biomass of dwarf Rhizophora mangle trees. Total aboveground biomass and their components were best estimated as a power function of the crown area times number of prop roots as an independent variable (Y = B ?? Xa??0.5083). The allometric equation for each tree component was highly significant (p<0.0001), with all r2 values greater than 0.90. The allometric relationship was used to estimate total aboveground biomass that ranged from 7.9 to 23.2 ton haa??1. Rhizophora mangle contributed 85% of total standing crop biomass. Conocarpus erectus, Laguncularia racemosa, and Avicennia germinans contributed the remaining biomass. Average aboveground biomass allocation was 69% for prop roots, 25% for stem and branches, and 6% for leaves. This aboveground biomass partitioning pattern, which gives a major role to prop roots that have the potential to produce an extensive root system, may be an important biological strategy in response to low phosphorus availability and relatively reduced soils that characterize mangrove forests in South Florida.

Florida

Effects of thinning on drought vulnerability and climate response in north temperate forest ecosystems

Reducing tree densities through silvicultural thinning has been widely advocated as a strategy for enhancing resistance and resilience to drought, yet few empirical evaluations of this approach exist. We examined detailed dendrochronological data from a long-term (>50 yrs) replicated thinning experiment to determine if density reductions conferred greater resistance and/or resilience to droughts, assessed by the magnitude of stand-level growth reductions. Our results suggest that thinning generally enhanced drought resistance and resilience; however, this relationship showed a pronounced reversal over time in stands maintained at lower tree densities. Specifically, lower-density stands exhibited greater resistance and resilience at younger ages (49 years), yet exhibited lower resistance and resilience at older ages (76 years), relative to higher-density stands. We attribute this reversal to significantly greater tree sizes attained within the lower-density stands through stand development, which in turn increased tree-level water demand during the later droughts. Results from response-function analyses indicate that thinning altered growth-climate relationships, such that higher-density stands were more sensitive to growing-season precipitation relative to lower-density stands. These results confirm the potential of density management to moderate drought impacts on growth, and they highlight the importance of accounting for stand structure when predicting climate-change impacts to forest systems.

Minnesota

River flow mass exponents with fractal channel networks and rainfall

An important problem in hydrologic science is understanding how river flow is influenced by rainfall properties and drainage basin characteristics. In this paper we consider one approach, the use of mass exponents, in examining the relation of river flow to rainfall and the channel network, which provides the primary conduit for transport of water to the outlet in a large basin. Mass exponents, which characterize the power-law behavior of moments as a function of scale, are ideally suited for defining scaling behavior of processes that exhibit a high degree of variability or intermittency. The main result in this paper is an expression relating the mass exponent of flow resulting from an instantaneous burst of rainfall to the mass exponents of spatial rainfall and that of the network width function. Spatial rainfall is modeled as a random multiplicative cascade and the channel network as a recursive replacement tree; these fractal models reproduce certain types of self-similar behavior seen in actual rainfall and networks. It is shown that under these modeling assumptions the scaling behavior of flow mirrors that of rainfall if rainfall is highly variable in space, and on the other hand flow mirrors the structure of the network if rainfall is not so highly variable. ?? 2001 Elsevier Science Ltd. All rights reserved.

Advances in Water Resources

Spatial variability and landscape controls of near-surface permafrost within the Alaskan Yukon River Basin

The distribution of permafrost is important to understand because of permafrost's influence on high-latitude ecosystem structure and functions. Moreover, near-surface (defined here as within 1&thinsp;m of the Earth's surface) permafrost is particularly susceptible to a warming climate and is generally poorly mapped at regional scales. Subsequently, our objectives were to (1) develop the first-known binary and probabilistic maps of near-surface permafrost distributions at a 30 m resolution in the Alaskan Yukon River Basin by employing decision tree models, field measurements, and remotely sensed and mapped biophysical data; (2) evaluate the relative contribution of 39 biophysical variables used in the models; and (3) assess the landscape-scale factors controlling spatial variations in permafrost extent. Areas estimated to be present and absent of near-surface permafrost occupy approximately 46% and 45% of the Alaskan Yukon River Basin, respectively; masked areas (e.g., water and developed) account for the remaining 9% of the landscape. Strong predictors of near-surface permafrost include climatic indices, land cover, topography, and Landsat 7 Enhanced Thematic Mapper Plus spectral information. Our quantitative modeling approach enabled us to generate regional near-surface permafrost maps and provide essential information for resource managers and modelers to better understand near-surface permafrost distribution and how it relates to environmental factors and conditions.

Alaska

Bryophytes and lichens: Small but indispensable forest dwellers

* What is a Bryophyte? * Bryophytes are the small green plants commonly known as mosses, liverworts and hornworts. Compared to plants, they have primitive tissues for conducting food and water, and they lack a protective outer surface to maintain water balance. Most bryophytes, because they lack tissues such as roots, obtain their water through direct surface contact with their environment. During dry weather they have the capacity to withstand complete dehydration. Bryophytes that are dry may appear dead but will regain normal function when moisture is available. Instead of producing seeds, bryophytes can either reproduce sexually by means of spores, or asexually when small pieces break off and grow into new individuals. * What is a Lichen? * Lichens are dual organisms consisting of a fungus and an alga or a cyanobacterium. The fungus provides the alga with structure, protection, nutrients, and water absorbed from the atmosphere and the substrate (e.g., soil, rotten logs, tree branches). In return, the alga provides carbohydrates from photosynthesis to the fungus. Algae from some lichens grow independently of the fungus, but in lichen form, the algae can inhabit more challenging environments than when growing alone. Most lichens can reproduce asexually. They either produce specialized propagules containing both partners, or parts of the lichen simply break, allowing both the fungus and the alga to disperse together. In some lichens, the fungal partner reproduces sexually by releasing spores, but the partner alga must be present in order for a lichen to reform.

Fact Sheet

Arthropods of Rose Atoll with special reference to ants and Pulvinaria Urbicola Scales (Hempitera Coccidae) on Pisonia Grandis trees

Rose Atoll, at the eastern end of the Samoan Archipelago, is a small but important refuge for seabirds, shorebirds, and sea turtles. While the vertebrate community is relatively well-studied, the terrestrial arthropod fauna, and its role in ecosystem function, are poorly known. Arthropods may be influencing the decline of Pisonia grandis, an ecologically important tree that once dominated the 6.6 ha of land on Rose Atoll. Reasons for the decline are not fully understood but a facultative relationship between two invasive arthropods, the soft scale Pulvinaria urbicola and ants, likely has contributed to tree death. The primary objectives of this study were to systematically survey the terrestrial arthropod fauna and identify ant species that tend scales on Pisonia. Using an array of standard arthropod collecting techniques, at least 73 species from 20 orders were identified, including nine ant species. Of the ants collected, only Tetramorium bicarinatum and T. simillimum were observed tending scales on Pisonia. No known natural enemies of Pulvinaria scales were found, suggesting little predation on scale populations. Treatment of Pisonia with the systemic insecticide imidacloprid failed to eliminate Pulvinaria scales, although short-term suppression apparently occurred. The arthropod fauna of Rose Atoll is dominated by exotic species that likely have a significant impact on the structure and function of the island&rsquo;s ecosystem.

Rose Atoll; America Samoa

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Nature Ecology & Evolution

Biogeography of fire regimes in western US conifer forests: A trait-based approach

Aim Functional traits are a critical link between species distributions and the ecosystem processes that structure those species’ niches. Concurrent increases in the availability of functional trait data and our ability to model species distributions present an opportunity to develop functional trait biogeography, i.e. the mapping of functional traits across space. Functional trait biogeography can improve process-based predictions about the resistance of certain species assemblages to changing environmental conditions across landscape scales. We illustrate this concept by developing the first trait-based, quantitative ranking of fire resistance (adult tree survival) in North American conifer species, and mapping that fire resistance across space. Location and Time period Western Continental United States, present-day. Major taxa studied 29 common conifer tree species. Methods We compiled six traits for each species: three relating to tree morphology and three relating to litter flammability. We combined these traits into a single fire resistance score, and used community-weighted averaging to estimate the fire resistance scores of different forest communities, using interpolated species distribution and relative abundance data. Results Species associated with historically frequent fire have high fire resistance scores (e.g., Pinus ponderosa), reflected by thick bark, tall crowns, and flammable litter. Species associated with subalpine or arid conditions have low fire resistance scores (e.g., Picea engelmannii and Pinus edulis), reflected by thin bark, short stature, poor self-pruning and low litter flammability. A map of forest community fire resistance across the western US reveals agreement with independent assessments of historical fire regimes, while also identifying areas where community-wide species traits may be mismatched with historical fire regimes. Main conclusions Quantifying the functional traits that confer resistance to tree-killing fire provides a direct link between ecosystem disturbance and community resistance. Understanding this link is critical to evaluating long-term resilience of different forest types under dynamic fire regimes. Our work represents the first known spatial representation of fire-resistance traits at a regional scale, and as such provides a link between functional traits and biogeography relevant to a critical ecosystem process.

Global Ecology and Biogeography

Conservation genomics of an endangered montane amphibian reveals low population structure, low genomic diversity and selection pressure from disease

Wildlife diseases are a major global threat to biodiversity. Boreal toads ( Anaxyrus [Bufo] boreas ) are a state-endangered species in the southern Rocky Mountains of Colorado and New Mexico, and a species of concern in Wyoming, largely due to lethal skin infections caused by the amphibian chytrid fungus Batrachochytrium dendrobatidis ( Bd ). We performed conservation and landscape genomic analyses using single nucleotide polymorphisms from double-digest, restriction site-associated DNA sequencing in combination with the development of the first boreal toad (and first North American toad) reference genome to investigate population structure, genomic diversity, landscape connectivity and adaptive divergence. Genomic diversity ( π = 0.00034–0.00040) and effective population sizes ( N e = 8.9–38.4) were low, likely due to post-Pleistocene founder effects and Bd -related population crashes over the last three decades. Population structure was also low, likely due to formerly high connectivity among a higher density of geographically proximate populations. Boreal toad gene flow was facilitated by low precipitation, cold minimum temperatures, less tree canopy, low heat load and less urbanization. We found >8X more putatively adaptive loci related to Bd intensity than to all other environmental factors combined, and evidence for genes under selection related to immune response, heart development and regulation and skin function. These data suggest boreal toads in habitats with Bd have experienced stronger selection pressure from disease than from other, broad-scale environmental variations. These findings can be used by managers to conserve and recover the species through actions including reintroduction and supplementation of populations that have declined due to Bd .

Colorado