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Kueppers, Lara M.

Publications and source records attributed to Kueppers, Lara M..

Needle bacterial community structure across the species range of limber pine

Abstract Bacteria on and inside leaves can influence forest tree health and resilience. The distribution and limits of a tree species’ range can be influenced by various factors, with biological interactions among the most significant. We investigated the processes shaping the bacterial needle community across the species distribution of limber pine, a widespread Western conifer inhabiting a range of extreme habitats. We tested four hypotheses: (i) Needle community structure varies across sites, with site-specific factors more important to microbial assembly than host species selection; (ii) dispersal limitation structures foliar communities across the range of limber pine; (iii) the relative significance of dispersal and selection differs across sites in the tree species range; and (iv) needle age structures bacterial communities. We characterized needle communities from the needle surface and tissue of limber pine and co-occurring conifers across 16 sites in the limber pine distribution. Our findings confirmed that site characteristics shape the assembly of bacterial communities across the host species range and showed that these patterns are not driven by dispersal limitation. Furthermore, the strength of selection by the host varied by site, possibly due to differences in available microbes. Our study, by focusing on trees in their natural setting, reveals real needle bacterial dynamics in forests, which is key to understanding the balance between stochastic and deterministic processes in shaping forest tree-microbe interactions. Such understanding will be necessary to predict or manipulate these interactions to support forest ecosystem productivity or assist plant migration and adaptation in the face of global change.

59 BASIC BIOLOGICAL SCIENCES↗

Allometric relationships and trade‐offs in 11 common M editerranean‐climate grasses

Abstract Biomass allocation in plants is the foundation for understanding dynamics in ecosystem carbon balance, species competition, and plant–environment interactions. However, existing work on plant allometry has mainly focused on trees, with fewer studies having developed allometric equations for grasses. Grasses with different life histories can vary in their carbon investment by prioritizing the growth of specific organs to survive, outcompete co‐occurring plants, and ensure population persistence. Further, because grasses are important fuels for wildfire, the lack of grass allocation data adds uncertainty to process‐based models that relate plant physiology to wildfire dynamics. To fill this gap, we conducted a greenhouse experiment with 11 common California grasses varying in photosynthetic pathway and growth form. We measured plant sizes and harvested above‐ and belowground biomass throughout the life cycle of annual species, while for the establishment stage of perennial grasses to quantify allometric relationships for leaf, stem, and root biomass, as well as plant height and canopy area. We used basal diameter as a reference measure of plant size. Overall, basal diameter is the best predictor for leaf and stem biomass, height, and canopy area. Including height as another predictor can improve model accuracy in predicting leaf and stem biomass and canopy area. Fine root biomass is a function of leaf biomass alone. Species vary in their allometric relationships, with most variation occurring for plant height, canopy area, and stem biomass. We further explored potential trade‐offs in biomass allocation across species between leaf and fine root, leaf and stem, and allocation to reproduction. Consistent with our expectation, we found that fast‐growing plants allocated a greater fraction to reproduction. Additionally, plant height and specific leaf area negatively influenced the leaf‐to‐stem ratio. However, contrary to our hypothesis, there were no differences in root‐to‐leaf ratio between perennial and annual or C 4 and C 3 plants. Our study provides species‐specific and functional‐type‐specific allometry equations for both above‐ and belowground organs of 11 common California grass species, enabling nondestructive biomass assessment in California grasslands. These allometric relationships and trade‐offs in carbon allocation across species can improve ecosystem model predictions of grassland species interactions and environmental responses through differences in morphology.

54 ENVIRONMENTAL SCIENCES↗

Nutrient Dynamics in a Coupled Terrestrial Biosphere and Land Model (ELM-FATES-CNP)

We present a representation of nitrogen and phosphorus cycling in the Functionally Assembled Terrestrial Ecosystem Simulator, a demographic vegetation model within the Energy Exascale Earth System land model. This representation is modular, and designed to allow testing of multiple hypothetical approaches for carbon-nutrient coupling in plants. Novel model hypotheses introduced in this work include, (a) the controls on plant acquisition of aqueous mineralized nutrients in the soil and (b) fairly straight forward methods of allocating nutrients to specific plant organs and their losses through live plant turnover as well as litter fluxes generated through plant mortality. This combines the new with pre-existing hypotheses (such as nitrogen fixation and soil decomposition) into a system that can accommodate plant-soil dynamics for a large number of size- and functional-type-resolved plant cohorts within a time-since-disturbance-resolved ecosystem. Root uptake of nutrients is governed by fine root biomass, and plants vary in their fine root biomass allocation in order to balance carbon and nutrient limitations to growth. We test the sensitivity of the model to a wide range of parameter variations and structural representations, and in the context of observations at Barro Colorado Island, Panama. A key model prediction is that plants in the high-light-availability canopy positions allocate more carbon to fine roots than plants in low-light understory environments, given the widely different carbon versus nutrient constraints of these two niches within a given ecosystem. This model provides a basis for exploring carbon-nutrient coupling with vegetation demography within Earth system models.

54 ENVIRONMENTAL SCIENCES↗

w Mel replacement of dengue-competent mosquitoes is robust to near-term climate change

Rising temperatures are impacting the range and prevalence of mosquito-borne diseases. A promising biocontrol technology replaces wild mosquitoes with those carrying the virus-blocking Wolbachia bacterium. Because the most widely used strain, wMel, is adversely affected by heat stress, we examined how global warming may influence wMel-based replacement. We simulated interventions in two locations with successful field trials using Coupled Model Intercomparison Project Phase 5 climate projections and historical temperature records, integrating empirical data on wMel’s thermal sensitivity into a model of Aedes aegypti population dynamics to evaluate introgression and persistence over one year. We show that in Cairns, Australia, climatic futures necessitate operational adaptations for heatwaves exceeding two weeks. In Nha Trang, Vietnam, projected heatwaves of three weeks and longer eliminate wMel under the most stringent assumptions of that symbiont’s thermal limits. We conclude that this technology is generally robust to near-term (2030s) climate change. Accelerated warming may challenge this in the 2050s and beyond.

54 ENVIRONMENTAL SCIENCES↗

Carbon stored in live ponderosa pines in the Sierra Nevada will not return to pre-drought (2012) levels during the 21st century due to bark beetle outbreaks

Outbreaks of several bark beetle species can develop rapidly in response to drought and may result in large transfers of carbon (C) stored in live trees to C stored in dead trees (10s of Tg C yr -1 in the western U.S. alone), which over time will be released back to the atmosphere. The western pine beetle (WPB) outbreak incited by the 2012–2015 mega-drought in the Sierra Nevada, California, U.S., could portend more frequent and/or severe bark beetle outbreaks as the temperature warms and drought frequency and intensity increase in the future. However, changes in the frequency and/or severity (resultant levels of host tree mortality) of beetle outbreaks are difficult to predict as outbreaks are complex with non-linear and eruptive processes primarily driven by interactions among beetle populations, the demography of hosts and other tree species, and climate and weather. Using an insect phenology and tree defense model, we projected the future likelihood of WPB outbreaks in the Sierra Nevada with climate drivers from different Earth System Models. Our goal was to understand how host (ponderosa pine, PIPO) recovery and future warming and drought affect the frequency and severity of WPB outbreaks and their C consequences. Our projections suggested that by 2100 the C stored in live PIPO (mean: 1.98 kg C m -2 , 95% CI: 1.74–2.21 kg C m -2 ) will not return to levels that occurred before the 2012–2015 drought (2012: ~2.30 kg C m –2 ) due to future WPB outbreaks. However, differences in climate models indicate a wide range of possible WPB outbreak frequencies and severities. Our results suggest that total plot basal area is the most significant factor in the mortality rate of PIPO by WPB in any given year, followed by drought severity and temperature. High levels of host basal area, higher temperature, and extreme drought all contribute to the frequency and severity of future WPB outbreaks. While PIPO basal area may decline under increased drought and warming, limiting high-stand basal area (>60 m 2 ha -1 ) may reduce the severity of future WPB outbreaks in the Sierra Nevada.

54 ENVIRONMENTAL SCIENCES↗

Data from: "Moisture rivals temperature in limiting photosynthesis by trees establishing beyond their cold-edge range limit under ambient and warmed conditions"

This archive contains data files that were used to draw conclusions in “Moisture rivals temperature in limiting photosynthesis by trees establishing beyond their cold-edge range limit under ambient and warmed conditions”, by Moyes et al., 2015. All field research was completed in common garden plots set up as part of the Alpine Treeline Warming Experiment (ATWE) on Niwot Ridge, Colorado, USA.There are two main data file formats in this archive: comma-separated values (.csv), and Microsoft Excel (.xls and .xlsx). .xlsx files can be read using Microsoft Excel and Google Sheets, and .csv files can be read using any simple text editor program, such as TextEdit (Mac) and Notepad (Windows). This .pdf data user’s guide can be read using Adobe Acrobat Reader, or any other compatible software. Seedling photographs and their corresponding leaf area-processed images are available in .jpg/.JPG image format, and can be opened using Preview (Mac) and Photos (Windows). To provide additional spatial context, two types of geospatial files are also published in this data package: ESRI shapefiles (.shp) and .kml files. Shapefiles are compatible with any GIS software able to read the file type (such as QGIS or ESRI’s ArcGIS suite), and .kml files can be opened with Google Earth or Google Maps. Figures 3 and 4 in the publication contain data from Moyes et al. 2013. This publication is cited in the References section in this archive, and data files can be accessed via the Alpine Treeline Warming Experiment project portal on ESS-DIVE. ---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------- Climate change is altering plant species distributions globally, and warming is expected to promote uphill shifts in mountain trees. However, at many cold-edge range limits, such as alpine treelines in the western United States, tree establishment may be colimited by low temperature and low moisture, making recruitment patterns with warming difficult to predict.- We measured response functions linking carbon (C) assimilation and temperature- and moisture-related microclimatic factors for limber pine (Pinus flexilis) seedlings growing in a heating × watering experiment within and above the alpine treeline. We then extrapolated these response functions using observed microclimate conditions to estimate the net effects of warming and associated soil drying on C assimilation across an entire growing season.- Moisture and temperature limitations were each estimated to reduce potential growing season C gain from a theoretical upper limit by 15–30% (c. 50% combined). Warming above current treeline conditions provided relatively little benefit to modeled net assimilation, whereas assimilation was sensitive to either wetter or drier conditions.- Summer precipitation may be at least as important as temperature in constraining C gain by establishing subalpine trees at and above current alpine treelines as seasonally dry subalpine and alpine ecosystems continue to warm.

54 ENVIRONMENTAL SCIENCES↗

Linking soil phosphorus with forest litterfall resistance and resilience to cyclone disturbance: A pantropical meta-analysis

While tropical cyclone regimes are shifting with climate change, the mechanisms underpinning the resistance (ability to withstand disturbance-induced change) and resilience (capacity to return to pre-disturbance reference) of tropical forest litterfall to cyclones remain largely unexplored pantropically. Single-site studies in Australia and Hawaii suggest that litterfall on low-phosphorus (P) soils is more resistant and less resilient to cyclones. Here, we conducted a meta-analysis to investigate the pantropical importance of total soil P in mediating forest litterfall resistance and resilience to 22 tropical cyclones. We evaluated cyclone-induced and post-cyclone litterfall mass (g/m 2 /day), and P and nitrogen (N) fluxes (mg/m 2 /day) and concentrations (mg/g), all indicators of ecosystem function and essential for nutrient cycling. Across 73 case studies in Australia, Guadeloupe, Hawaii, Mexico, Puerto Rico, and Taiwan, total litterfall mass flux increased from ~2.5 ± 0.3 to 22.5 ± 3 g/m 2 /day due to cyclones, with large variation among studies. Litterfall P and N fluxes post-cyclone represented ~5% and 10% of the average annual fluxes, respectively. Post-cyclone leaf litterfall N and P concentrations were 21.6 ± 1.2% and 58.6 ± 2.3% higher than pre-cyclone means. Mixed-effects models determined that soil P negatively moderated the pantropical litterfall resistance to cyclones, with a 100 mg P/kg increase in soil P corresponding to a 32% to 38% decrease in resistance. Based on 33% of the resistance case studies, total litterfall mass flux reached pre-disturbance levels within one-year post-disturbance. A GAMM indicated that soil P, gale wind duration and time post-cyclone jointly moderate the short-term resilience of total litterfall, with the nature of the relationship between resilience and soil P contingent on time and wind duration. Across pantropical forests observed to date, our results indicate that litterfall resistance and resilience in the face of intensifying cyclones will be partially determined by total soil P.

59 BASIC BIOLOGICAL SCIENCES↗

Forest regeneration within Earth system models: current process representations and ways forward

Earth system models must predict forest responses to global change in order to simulate future global climate, hydrology, and ecosystem dynamics. These models are increasingly adopting vegetation demographic approaches that explicitly represent tree growth, mortality, and recruitment, enabling advances in the projection of forest vulnerability and resilience, as well as evaluation with field data. To date, simulation of regeneration processes has received far less attention than simulation of processes that affect growth and mortality, in spite of their critical role maintaining forest structure, facilitating turnover in forest composition over space and time, enabling recovery from disturbance, and regulating climate-driven range shifts. Here, our critical review of regeneration process representations within current Earth system vegetation demographic models reveals the need to improve parameter values and algorithms for reproductive allocation, dispersal, seed survival and germination, environmental filtering in the seedling layer, and tree regeneration strategies adapted to wind, fire, and anthropogenic disturbance regimes. These improvements require synthesis of existing data, specific field data-collection protocols, and novel model algorithms compatible with global-scale simulations. Vegetation demographic models offer the opportunity to more fully integrate ecological understanding into Earth system prediction; regeneration processes need to be a critical part of the effort.

54 ENVIRONMENTAL SCIENCES↗

Simulating environmentally-sensitive tree recruitment in vegetation demographic models

Vegetation demographic models (VDMs) endeavor to predict how global forests will respond to climate change. This requires simulating which trees, if any, are able to recruit under changing environmental conditions. We present a new recruitment scheme for VDMs in which functional-type-specific recruitment rates are sensitive to light, soil moisture and the productivity of reproductive trees. We evaluate the scheme by predicting tree recruitment for four tropical tree functional types under varying meteorology and canopy structure at Barro Colorado Island, Panama. We compare predictions to those of a current VDM, quantitative observations and ecological expectations. We find that the scheme improves the magnitude and rank order of recruitment rates among functional types and captures recruitment limitations in response to variable understory light, soil moisture and precipitation regimes. Furthermore, our results indicate that adopting this framework will improve VDM capacity to predict functional-type-specific tree recruitment in response to climate change, thereby improving predictions of future forest distribution, composition and function.

59 BASIC BIOLOGICAL SCIENCES↗

Improving sustainable tropical forest management with voluntary carbon markets

Due to a rapidly changing climate, voluntary carbon markets are gaining momentum and should be leveraged to improve and expand tropical sustainable forest management plans, limiting carbon emissions and enhancing critical carbon sinks. By sequestering more carbon than any other terrestrial ecosystem — ~1 Pg C yr –1 — tropical forests provide crucial natural climate solutions and opportunities in the evolving voluntary carbon market. Here, we argue that some issues with the current sustainable management of tropical forests can be addressed using carbon-focused sustainable forest management (SFM + C) to leverage financial resources for tropical forest carbon storage and sequestration. We suggest an extended harvest cycle in SFM + C and calculate an associated potential increase in aboveground carbon stocks of commercial timber of 1.26 Mg C ha –1 after each cycle in the Brazilian Amazon. The additional carbon storage due to a longer harvest cycle can generate carbon credits worth 152.6 (SD 9.2) US dollars per hectare in 40 years. Considering an average cost of 180 BRL per m3 of commercial timber delivered to the sawmill, an SFM + C plan with a 40-year cycle could generate 28.7% (SD 2.5) more profit than 35-year cycles by combining timber and carbon revenues. A robust carbon price could incentivize the further extension of harvest cycles, providing a monetary return that offsets the opportunity cost intrinsic to harvesting under longer cycles. Lastly, we highlight research needs to support tropical SFM + C, which can be part of a global collective effort to limit global warming to below 2 °C above pre-industrial levels.

54 ENVIRONMENTAL SCIENCES↗

Data from: "Lab and Field Warming Similarly Advance Germination Date and Limit Germination Rate for High and Low Elevation Provenances of Two Widespread Subalpine Conifers"

This archive contains data used to draw conclusions in “Lab and Field Warming Similarly Advance Germination Date and Limit Germination Rate for High and Low Elevation Provenances of Two Widespread Subalpine Conifers”, by Kueppers et al. 2017. Cone collection and field experiments were conducted on Niwot Ridge, Colorado, with the latter completed within the Alpine Treeline Warming Experiment (ATWE). Laboratory germination experiments were conducted at the University of California Merced. Geospatial files are also provided for additional geographical context.There are four data file formats in this package: comma-separated values (.csv), Microsoft Excel (.xlsx), keyhole markup language (.kml), and ESRI shapefile (.shp). The latter two are geospatial file formats. .csv files can be opened using any simple text editor program such as TextEdit (Mac) and Notepad (Windows), and .xlsx files can be opened using Microsoft Excel or Google Sheets. The .kml file contains coordinate data, and is compatible with Google Earth and Google Maps. ESRI shapefiles (.shp) are compatible with any geospatial software able to read the file type, such as QGIS and ESRI’s ArcGIS Suite. This data user’s guide is available in as a .pdf, and can be opened using software such as Adobe Acrobat Reader.-----------------------------------------------------------------------------------------------------------------------------------Accurately predicting upslope shifts in subalpine tree ranges with warming requires understanding how future forest populations will be affected by climate change, as these are the seed sources for new tree line and alpine populations. Early life history stages are particularly sensitive to climate and are also influenced by genetic variation among populations. We tested the climate sensitivity of germination and initial development for two widely distributed subalpine conifers, using controlled-environment growth chambers with one temperature regime from subalpine forest in the Colorado Rocky Mountains and one 5 °C warmer, and two soil moisture levels. We tracked germination rate and timing, rate of seedling development, and seedling morphology for two seed provenances separated by ~300 m elevation. Warming advanced germination timing and initial seedling development by a total of ~2 weeks, advances comparable to mean differences between provenances. Advances were similar for both provenances and species; however, warming reduced the overall germination rate, as did low soil moisture, only for Picea engelmannii. A three-year field warming and watering experiment planted with the same species and provenances yielded responses qualitatively consistent with the lab trials. Together these experiments indicate that in a warmer, drier climate, P. engelmannii germination, and thus regeneration, could decline, which could lead to declining subalpine forest populations, while Pinus flexilis forest populations could remain robust as a seed source for upslope range shifts.

54 ENVIRONMENTAL SCIENCES↗

Data from: "Warming and the dependence of limber pine (Pinus flexilis) establishment on summer soil moisture within and above its current elevation range"

This data package contains data that were used for analysis in “Warming and the dependence of limber pine (Pinus flexilis) establishment on summer soil moisture within and above its current elevation range”, by Moyes et al. 2013. All data collection and field research were completed on Niwot Ridge, Colorado, USA.This data package contains nine comma-separated-values (.csv) files, one text (.txt) file, and two zipped seedling folders that were used for leaf area analysis. One zipped folder contains 468 .jpg photographs of seedlings, and the second contains 468 corresponding Image J-processed .jpg images that include silhouette leaf area values. .csv and .txt files can be opened using any compatible simple text-editor software such as TextEdit (Mac) and Notepad (Windows); .csv’s can also be opened using R and Microsoft Excel. Image files can be opened using Preview (Mac) and Photos (Windows). In addition, there are a total of 31 Microsoft Excel files: three .xlsx files, and 28 raw Li-Cor output .xls files. This data user’s guide is available in .pdf format, and can be opened using Adobe Acrobat Reader, or any other compatible file viewing software. Geospatial data showing field site locations are also included in the archive for use and reference. There are two geospatial formats in this archive: ESRI shapefiles (.shp) and keyhole markup-language (.kml) files. Both file types contain bounding box information, with the former being polygons, and the latter containing corner coordinates for each site. ESRI shapefiles can be opened using any geospatial software compatible with the file type (such as ESRI’s ArcGIS suite and QGIS), and .kml files are compatible with Google Earth and Google Maps.--------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------Continued changes in climate are projected to alter the geographic distributions of plant species, in part by affecting where individuals can establish from seed. We tested the hypothesis that warming promotes uphill redistribution of subalpine tree populations by reducing cold limitation at high elevation and enhancing drought stress at low elevation. We seeded limber pine (Pinus flexilis) into plots with combinations of infrared heating and water addition treatments, at sites positioned in lower subalpine forest, the treeline ecotone, and alpine tundra. In 2010, first-year seedlings were assessed for physiological performance and survival over the snow-free growing season. Seedlings emerged in midsummer, about 5–8 weeks after snowmelt. Low temperature was not observed to limit seedling photosynthesis or respiration between emergence and October, and thus experimental warming did not appear to reduce cold limitation at high elevation. Instead, gas exchange and water potential from all sites indicated a prevailing effect of summer moisture stress on photosynthesis and carbon balance. Infrared heaters raised soil growing degree days (base 5 °C, p < 0.001) and August–September mean soil temperature (p < 0.001). Despite marked differences in vegetation cover and meteorological conditions across sites, volumetric soil moisture content (θ) at 5–10 cm below 0.16 and 0.08 m^3 m^(−3) consistently corresponded with moderate and severe indications of drought stress in midday stem water potential, stomatal conductance, photosynthesis, and respiration. Seedling survival was greater in watered plots than in heated plots (p = 0.01), and negatively related to soil growing degree days and duration of exposure to θ < 0.08 m^3 m^(−3) in a stepwise linear regression model (p < 0.0001). We concluded that seasonal moisture stress and high soil surface temperature imposed a strong limitation to limber pine seedling establishment across a broad elevation gradient, including at treeline, and that these limitations are likely to be enhanced by further climate warming.--------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------Maintenance log:Amended file name error in Data User's Guide May 19 2022

54 ENVIRONMENTAL SCIENCES↗

Warming increased bark beetle-induced tree mortality by 30% during an extreme drought in California

Quantifying the responses of forest disturbances to climate warming is critical to our understanding of carbon cycles and energy balances of the Earth system. The impact of warming on bark beetle outbreaks is complex as multiple drivers of these events may respond differently to warming. Using a novel model of bark beetle biology and host tree interactions, we assessed how contemporary warming affected western pine beetle (Dendroctonus brevicomis) populations and mortality of its host, ponderosa pine (Pinus ponderosa), during an extreme drought in the Sierra Nevada, California, United States. When compared with the field data, our model captured the western pine beetle flight timing and rates of ponderosa pine mortality observed during the drought. In assessing the influence of temperature on western pine beetles, we found that contemporary warming increased the development rate of the western pine beetle and decreased the overwinter mortality rate of western pine beetle larvae leading to increased population growth during periods of lowered tree defense. We attribute a 29.9% (95% CI: 29.4%–30.2%) increase in ponderosa pine mortality during drought directly to increases in western pine beetle voltinism (i.e., associated with increased development rates of western pine beetle) and, to a much lesser extent, reductions in overwintering mortality. These findings, along with other studies, suggest each degree (°C) increase in temperature may have increased the number of ponderosa pine killed by upwards of 35%–40% °C-1 if the effects of compromised tree defenses (15%–20%) and increased western pine beetle populations (20%) are additive. Due to the warming ability to considerably increase mortality through the mechanism of bark beetle populations, models need to consider climate's influence on both host tree stress and the bark beetle population dynamics when determining future levels of tree mortality.

59 BASIC BIOLOGICAL SCIENCES↗