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Bradley J. Butterfield

Publications and source records attributed to Bradley J. Butterfield.

At least 19 recordsLinked to original sources

At the edge of change: Native Phragmites persistence and nonnative arrival along the dam-regulated Colorado River

Premise The southwestern United States is at the forefront of nonnative Phragmites australis subsp. australis expansion. Native Phragmites australis subsp. americanus , likely in decline in North America, is ecologically and culturally important and of conservation interest. In the greater Grand Canyon region, the Glen Canyon Dam led to loss of habitat upstream and increases in Phragmites downstream. Recent increases correspond with changes to dam operations and the arrival of nonnative Phragmites . Determining the native status of Phragmites stands along the Colorado River and around Lake Powell can inform complex management choices and restoration planning. Methods Phragmites subspecies at 83 locations from southern Utah through northern Arizona, United States were genetically tested using real-time PCR. Genetic diversity and structure were assessed using microsatellite loci for 13 native Phragmites stands. Morphological traits of southwestern native Phragmites were compared to those of nonnative Phragmites . Results Nonnative Phragmites was identified at four of the 83 locations. Native Phragmites in the study area is genetically diverse with little regional genetic structure, but substantial genetic differentiation among sites. Southwestern native Phragmites is morphologically variable, but characteristics can be used together to distinguish it from nonnative Phragmites . Conclusions The greater Grand Canyon and Glen Canyon regions support a large, genetically diverse native Phragmites population, which is potentially threatened by nonnative Phragmites and changes to river management. Determining conditions that would lead to nonnative Phragmites invasion of desert wetlands will facilitate discussions about the risks and benefits of Phragmites management in these unique ecosystems.

Arizona, Nevada, Utah

Functional traits can explain coordinated seedling responses to drought and freezing stress

Limited plant establishment creates demographic bottlenecks that can alter vegetation trajectories in changing environments, yet it is unclear how common eco-evolutionary indicators of plant strategy shape early response to key stressors. We evaluated how early functional traits, phylogeny, and adult climate niche jointly explain seedling drought and freezing tolerance across 49 semi-arid rangeland species. The best models explained 36% of variation in freezing tolerance (traits only) and 45% of variation in drought tolerance (traits, phylogeny, and climate), identifying both shared and unique indicators of different stress responses. Drought tolerance was explained by rapid root elongation and large seed size, while phylogeny captured additional tolerance within the grass family. Seedling freezing tolerance was linked to rapid root elongation and colder thresholds for root growth, though traits shared some explanatory power with adult climate niche (cold temperature distribution). While stress resistance can be shaped by phylogeny and adaptations to long-term climate regimes, our results elucidate early traits with clearer direct links to establishment capacity under drought and freezing stress, across species. Additional research is needed to understand how variation in stress response strategies across multiple demographic stages (e.g., seeds, seedlings, mature plant) and ecological scales (e.g., within and across species' climate distributions) could affect vegetation distributions and trajectories in changing environments.

Colorado Plateau, Great Basin, western United Stat

Aridity reduces lag times between aquatic and terrestrial dry-down among watersheds and across years in the northwest US

Landscapes encompass both aquatic and terrestrial ecosystems that experience the same climate but may respond to climate in divergent ways. For example, the time lag between seasonal dry-down of terrestrial soil moisture and decline in streamflow has important implications for species and ecosystem processes across the aquatic–terrestrial interface. How these lags between aquatic and terrestrial hydrology vary with climate and spatial location within watersheds remains largely unexplored. Here, we examine seasonal patterns of aquatic–terrestrial dry-down across seven watersheds in the northwestern USA, spanning a wide range of aridity. We compared daily streamflow data from USGS gages at watershed outlets with simulated daily soil moisture (1979–2020) from multiple locations within each watershed. In all watersheds, annual dry cycles progressed sequentially through the following features: evapotranspiration, precipitation, shallow soil moisture, deep soil moisture, and finally streamflow. Seasonal streamflow minima lagged behind soil moisture minima for shorter durations in more arid watersheds and drier years. Within watersheds, lag times varied spatially due to interactions between elevation and aridity, with short lags in low-elevation soils near streams in arid watersheds and longer lags in less arid watersheds. Collectively, these results indicate shorter lags between seasonal aquatic and terrestrial dry periods in drier watersheds and years, and show that these tighter linkages are spatially aggregated in drier watersheds. The co-occurrence of seasonally dry conditions in both aquatic and terrestrial systems under increasing aridification is likely to intensify stressors on ecosystems and services. Recognizing these patterns may be critical for predicting ecosystem vulnerabilities and informing adaptation strategies to mitigate the impacts of seasonally dry conditions.

Montana, Oregon, Wyoming

Daily fluctuating flows affect riparian plant species distributions from local to regional scales

Aims The number of hydropower dams has grown globally over recent decades, with significant impacts on downstream riparian plant communities. Many of these dams generate daily fluctuating flows known as hydropeaking to meet sub-daily variation in energy demands. Hydropeaking can significantly impact riparian plant communities, with obligate riparian species tending to experience the greatest negative effects on habitat suitability. Whether this pattern holds in arid biomes where daily soil moisture enhancements could benefit some plants is an open question. Location Colorado River, Grand Canyon, Western USA. Methods We used occurrence records to model species responses to variation in daily flow fluctuations across 32 689 river segments in the Western United States. We then applied estimates of hydropeaking responses derived from those models to understanding the abundance and fine scale hydrologic niches of riparian plant species in the Colorado River ecosystem downstream of Glen Canyon Dam, which has experienced vegetation expansion attributed to river regulation, including hydropeaking that began in 1964. Results At the regional scale, species with greater wetland dependence exhibited increasingly negative responses to hydropeaking across 1 496 species, consistent with previous studies at smaller scales. At the local scale of the Colorado River, we found that species inhabiting near-channel habitat characterized by daily inundation and exposure had positive modeled responses to hydropeaking, consistent with a long history of selection for species tolerant of hydropeaking. In contrast, species inhabiting the zone immediately above peak daily river stage had negative modeled responses to hydropeaking, suggesting that they are being excluded from otherwise suitable habitat nearer the channel. Conclusions These results demonstrate that hydropeaking can impact species distributions from local to regional scales by excluding obligate wetland species and reducing habitat suitability for some facultative wetland species. These results from an arid river system are consistent with those reported from other biomes.

Arizona

Hotter temperatures alter riparian plant outcomes under regulated river conditions

Climate change and river regulation alter environmental controls on riparian plant occurrence and cover worldwide. Simultaneous changes to river flow and air temperature could result in unanticipated plant responses to novel environmental conditions. Increasing temperature could alter riparian plant response to hydrology and other factors, while river regulation may exacerbate environmental stress through novel flows like those resulting from power generation. Further, plant establishment and growth may require differing conditions, which may be decoupled by novel conditions. Using a large dataset that spans a natural 5°C mean annual temperature (MAT) gradient and a Bayesian model that integrates plant occurrence and cover, we address four questions: (1) Does hotter MAT modify plant response to hydrology, substrate composition, topography, and cover of co-occurring plant species? (2) Does the timing of hydropower tides benefit some species over others? (3) Does dam-induced erosion hinder riparian species more than upland species? (4) Do occurrence and cover respond to different environmental variables, allowing for decoupling of life history processes? We addressed these questions with data collected along 364 km of the Colorado River downstream of Glen Canyon Dam, Arizona, United States of America. Occurrence and cover class were recorded in >10,000 plots from 2016 to 2020, along with environmental covariates that repeat across the climate gradient. For 36 species, plant occurrence and cover were modeled with respect to MAT, hydrology, substrate, topography, other plant cover, and their interactions with MAT. There were four key results. (1) Increasing MAT will not only directly influence plants but will mediate their responses to the environment, including greater dependence on stable water supplies. (2) The timing of hydropower tides shapes plant community composition. (3) Dam-related erosion has an outsized effect on riparian species, which could lead to a loss of regionally unique plant species. (4) For all species, the most important covariates driving occurrence differed from those for cover, suggesting the potential for these life stages to be decoupled. Not only will climate change and river regulation independently alter plant distributions, interactions among hotter temperature, dam-controlled flow patterns, and limited fine sediments will determine which species flourish or perish under future conditions.

Arizona

Most pinyon-juniper woodland species distributions are projected to shrink rather than shift under climate change

Pinyon–juniper (PJ) woodlands are among the most widespread ecosystems in rangelands of western North America, supporting diverse wildlife habitat, recreation, grazing, and cultural/spiritual enrichment. Anticipating future distribution shifts under changing climate will be critical to climate adaptation and conservation efforts in these ecosystems. Here, we evaluate drivers of PJ tree species’ distributions and project changes in response to future climate change. We developed species distribution models with dryland-focused predictors to project environmental suitability changes across the entirety of three pinyon and six juniper species ranges. We identify areas of robust suitability change by combining suitability projections from multiple emissions scenarios and time periods. PJ species’ suitabilities respond to many temperature and moisture covariates expected to change in the future. Projected responses among PJ species are highly variable, ranging from modest declines with concurrent gains for overall little net change to wide-ranging declines with no gains for overall range contractions. Environmental suitability is projected to decline broadly across the arid United States Southwest and remain relatively stable across the northern Great Basin and Colorado Plateau. Our results suggest unique responses of PJ species to future climate change. We found that species were projected to experience more losses than gains in suitability, for overall range shrinks rather than shifts. Land managers have the capacity to increase woodland resilience to drought, and our results can inform rangeland-wide management planning and conservation efforts in PJ woodlands.

Rangeland Ecology & Management

U.S. Geological Survey Grand Canyon Monitoring and Research Center: Proceedings of the fiscal year 2023 annual reporting meeting to the Glen Canyon Dam Adaptive Management Program

This proceedings is prepared for the USBR and Glen Canyon Dam Adaptive Management Program (GCDAMP) to account for work conducted and products delivered in FY 2023 by SBSC's Grand Canyon Monitoring and Research Center (GCMRC) and to inform the Technical Work Group of science conducted by GCMRC and its cooperators in support of the GCDAMP. It includes a summary of accomplishments, modifications to work plans, and results related to projects included in GCMRC’s FY 2021-23 Triennial Work Plan. This work was done to support the 11 resource goals identified in the Glen Canyon Dam Long-Term Experimental and Management Plan (LTEMP) Environmental Impact Statement and Record of Decision.

Arizona, Nevada, Utah

Modeling the impacts of Glen Canyon Dam operations on Colorado River resources

At the time of this report, the Bureau of Reclamation (Reclamation) is writing two supplemental Environmental Impact Statements (sEIS ) and a new Environmental Impact Statement (EIS) that will analyze the effects of changing water flow out of Glen Canyon Dam (GCD) (U.S. Department of Interior, 2024). These actions have the potential to affect downstream resources, including threatened and endangered species, in the Grand Canyon. This report covers modeling support provided on the two sEIS by the USGS Grand Canyon Monitoring and Research Center (GCMRC; U.S. Geological Survey, Southwest Biological Science Center). The first sEIS (Interim Guidelines sEIS) modifies Reclamation’s 2007 Colorado River Interim Guidelines for Lower Basin Shortages and Coordinated Operations for Lake Powell and Lake Mead (USBR, 2007) that determines annual water releases from GCD based on inflow to Lake Powell, the power generating requirements of GCD, and relative lake levels of Lake Powell and Lake Mead. Drought conditions have lowered the level of Lake Powell and may require GCD to release less water than was analyzed in the Interim EIS (7-million-acre feet/year). The effect of less water released, as well as lower lake levels and associated water quality concerns, on downstream resources was not analyzed in the 2007 Interim Guidelines EIS. Reclamation requested GCMRC support to provide models predicting the effects to resources of water releases lower than 7M acre feet/year, including models predicting effects to threatened and endangered species for use in a Biological Assessment.

Report

Divergent physiological responses of hydric and mesic riparian plant species to a Colorado River experimental flow

Riparian plant species can differ in their responses to streamflow variation in ways that strongly influence the composition and functioning of riparian plant communities. Quantifying these differences and the potential asymmetry of responses to low- versus high-flow phases of stream fluctuations is important for predicting and managing vegetation responses to variation in flow regimes. We measured the physiological response of two riparian plant species with different habitat preferences to an experimental flow that progressed from a low-flow to high-flow phase. Schedonorous arundinaceus , a hydric grass growing in near-channel habitat, exhibited significant and biologically substantial declines in stem water potential (SWP) during the low-flow phase of the experiment and a saturating increase in leaf relative water content (LRWC) during the high-flow phase. These patterns are consistent with cavitation risk in response to low-flow anomalies and saturating responses to high-flow anomalies in this hydric species. Pluchea sericea , a mesic shrub growing at intermediate elevations above the channel, exhibited a decrease in LRWC during the low-flow phase and an increase in SWP during the high-flow phase. These patterns are consistent with protection from stem cavitation risk during drought through leaf dehydration and opportunistic increases in water status in response to high-flow anomalies in this mesic species. The asymmetrical responses of both species to low- versus high-flow phases demonstrate unique physiological responses to flow anomalies of contrasting directions attributable to species habitat preferences and functional strategies that can be used by managers to predict non-linear vegetation responses to flow variation.

Plant Ecology

Inundation tolerance, rather than drought tolerance, predicts riparian plant distributions along a local hydrologic gradient

Riparian vegetation varies along hydrologic gradients, along which inundation and drought tend to be inversely correlated. Differentiating effects of inundation and drought on plant distributions is critical for predicting impacts of changes to baseflows and designing flow patterns to achieve vegetation objectives in regulated river systems. To this end, we conducted a greenhouse experiment where we decreased, increased, or maintained constant water levels experienced by a suite of riparian plant species. We related changes in new root growth and stomatal conductance under experimental conditions to species hydrologic niches in the field, specifically the median elevation at which they occur above the channel, along the regulated Colorado River in Grand Canyon. We found a significant negative relationship between root growth response to experimental inundation with increasing elevation above the channel in the field, and a negative response of stomatal conductance to inundation among the most xeric-adapted species. Drought responses were idiosyncratic with respect to hydrologic niche, and instead seemed to vary in relation to clonality and rooting depth. Several Salicaceae tree species that are uncommon along regulated rivers exhibited consistently negative responses to both drought and inundation relative to other species, which may explain their rarity. The results of this study suggest that riparian plant distributions along hydrologic gradients have been shaped primarily by current and past levels of inundation. However, future anticipated declines in the water table are likely to produce species-specific responses based on drought tolerance that may in part be predicted from the results of this experiment.

Wetlands

Do seeding and seedling planting result in similar restored plant communities?

Aims Restoration practitioners often face a tradeoff between low cost but risky seeding vs expensive but more reliable seedling planting to meet revegetation goals. Knowing under what environmental and management conditions direct seeding vs seedling planting benefit different species could improve restoration practice. Methods We compared seed emergence to planted-seedling survival among perennial herbaceous species commonly used in restoration across eight experimental restoration sites on the Colorado Plateau, USA. We used linear models to assess relationships between emergence and survival among species, and to assess the effects of site climate and seeding pre-treatments on those relationships. Results We found that among species, emergence was positively correlated with survival in the cooler sites, meaning that species with high emergence also had high survival and vice versa, but had no relationship in the hottest sites. Furthermore, pre-treatments to enhance soil moisture in seeded plots, specifically microtopography (pits) and mulch, also resulted in positive relationships between emergence and survival among species, while seeding without additional soil pre-treatments did not. Seedling planting cost 14 times as much as direct seeding alone, dropping to nine times as much when pre-treatments were combined with seeding. Conclusions Taken together, these results suggest that investments in seedling planting at hotter dryland sites, or in creating microtopography or mulching prior to seeding across sites, are likely to promote establishment success compared to simple seeding methods in degraded dryland ecosystems. These findings also identify opportunities for hybrid seeding and planting approaches that balance tradeoffs between risk and cost, respectively.

Applied Vegetation Science

Assessment of riparian vegetation patterns and change downstream from Glen Canyon Dam from 2014 to 2019

Changes in riparian vegetation cover and composition occur in relation to flow regime, geomorphic template, and climate, and can have cascading effects on aquatic and terrestrial ecosystems. Tracking such changes over time is therefore an important part of monitoring the condition and trajectory of riparian ecosystems. Maintaining diverse, self-sustaining riparian vegetation comprised of mostly native species is identified in the Glen Canyon Dam Long-Term Experimental and Management Plan as a key resource objective for the section of the Colorado River between Glen Canyon Dam and Lake Mead. The U.S. Geological Survey Grand Canyon Monitoring and Research Center implemented an annual monitoring program in 2014 to assess the status and trends of riparian vegetation along this section of river, particularly as they relate to flow regime. In this report, we summarize plant species composition and cover data collected under the annual monitoring program from 2014 to 2019, with special consideration given to the hydrologic position, associated geomorphic feature class, local climate patterns, native and nonnative species, and floristic region for key vegetation metrics and species. We divided the study area into four river segments (referred to as Glen Canyon, Marble Canyon, eastern Grand Canyon, and western Grand Canyon) on the basis of geography and floristic composition and calculated each recorded plant species’ relative frequency and foliar cover by river segment. These data were then used to evaluate species composition relationships among river segments, hydrologic zones, geomorphic features, and sampling years through ordination analysis. Temporal trends in our focal resource objectives—species richness, total foliar cover, proportion of native to nonnative species richness, proportion of native to nonnative species cover, Tamarix cover, Pluchea sericea cover, and Baccharis species cover—were assessed using mixed-effects models. Four patterns related to species composition emerged: (1) species composition of fixed-site sandbars differed from that of randomly selected sites (including randomly selected sandbars), (2) species composition of Glen Canyon sites differed from that of other previously identified floristic regions, (3) species composition differed across hydrologic zones related to dam operations, and (4) species composition within river segments did not change across years. For temporal patterns, four main findings emerged: (1) trends differed between fixed-sites and randomly selected sites; (2) although few directional changes were observed from 2014 to 2019, Baccharis species cover increased at randomly selected sites in areas influenced by daily water fluctuations; (3) native species cover and richness were greater than nonnative species cover and richness across all hydrologic zones; and (4) the temporal trend metrics used here can be used across floristic groups, enabling assessment of the Colorado River ecosystem as a whole. In addition to these findings, lists of recorded plant species are included as appendixes. The variations and patterns in vegetation status and trends presented in this report can be used as a baseline against which future monitoring can be compared.

Arizona

Plant water-use strategies predict restoration success across degraded drylands

Plant strategies for coping with water limitation are likely to mediate restoration outcomes in degraded dryland ecosystems. Trade-offs in traits related to water acquisition and use can intensify in more arid environments, making their effects on dryland restoration success even more salient. However, isolating the effects of drought responses from those of other environmental factors, as well as identifying the specific drought resistance traits that influence restoration success, can be difficult. In the present study, we couple a controlled dry-down experiment with a cross-site restoration field trial of out-planted seedlings (RestoreNet) using a suite of dryland herbaceous plant species from the same seed sources. We quantified interspecific variation in physiological responses to drought, specifically reductions in stomatal conductance ( g s ) and stem water potential (SWP), by comparing well-watered control plants to those experiencing decreasing soil moisture. Drought responses of SWP and g s varied independently among species, but both were related to survival in the cross-site restoration field trial when effect sizes were aggregated across all sites. Responses were consistent with acquisitive water-use strategies resulting in greater success, where species with greater declines in SWP or weaker declines in g s under drought had greater survival. The correlation between SWP drought response and survival also intensified in sites with lower accumulated precipitation following restoration. Differences among functional groups revealed two different paths to restoration success: forbs that maintain high g s and narrow safety margins to maximize exploitation of moisture pulses before going into drought dormancy, or C 4 grasses that maintain efficient water uptake in drying soils while risking cavitation. C 3 grass species varied between these two strategies. Synthesis and applications . Taken together, the results of this study and others conducted at RestoreNet sites indicate that while a diversity of physiological responses to drought may exist in dryland plant communities, successfully restoring herbaceous species through out-planting in degraded conditions is likely to be achieved with species that maximize water uptake via one of two strategies, with tolerance of low SWP being particularly important in the most arid settings.

Journal of Applied Ecology

Soil surface treatments and precipitation timing determine seedling development across southwestern US restoration sites

Restoration in dryland ecosystems often has poor success due to low and variable water availability, degraded soil conditions, and slow plant community recovery rates. Restoration treatments can mitigate these constraints but, because treatments and subsequent monitoring are typically limited in space and time, our understanding of their applicability across broader environmental gradients remains limited. To address this limitation, we implemented and monitored a standardized set of seeding and soil surface treatments (pits, mulch, and ConMod artificial nurse plants) designed to enhance soil moisture and seedling establishment across RestoreNet, a growing network of 21 diverse dryland restoration sites in the southwestern USA over 3 years. Generally, we found that the timing of precipitation relative to seeding and the use of soil surface treatments were more important in determining seeded species emergence, survival, and growth than site-specific characteristics. Using soil surface treatments in tandem with seeding promoted up to 3× greater seedling emergence densities compared with seeding alone. The positive effect of soil surface treatments became more prominent with increased cumulative precipitation since seeding. The seed mix type with species currently found within or near a site and adapted to the historical climate promoted greater seedling emergence densities compared with the seed mix type with species from warmer, drier conditions expected to perform well under climate change. Seed mix and soil surface treatments had a diminishing effect as plants developed beyond the first season of establishment. However, we found strong effects of the initial period seeded and of the precipitation leading up to each monitoring date on seedling survival over time, especially for annual and perennial forbs. The presence of exotic species exerted a negative influence on seedling survival and growth, but not initial emergence. Our findings suggest that seeded species recruitment across drylands can generally be promoted, regardless of location, by (1) incorporation of soil surface treatments, (2) employment of near-term seasonal climate forecasts, (3) suppression of exotic species, and (4) seeding at multiple times. Taken together, these results point to a multifaceted approach to ameliorate harsh environmental conditions for improved seeding success in drylands, both now and under expected aridification.

Ecological Applications

Provenance, genotype, and flooding influence growth and resource acquisition characteristics in a clonal, riparian shrub

Premise Riparian plants can exhibit intraspecific phenotypic variability across the landscape related to temperature and flooding gradients. Phenotypes that vary across a climate gradient are often partly genetically determined and may differ in their response to inundation. Changes to inundation patterns across a climate gradient could thus result in site-specific inundation responses. Phenotypic variability is more often studied in riparian trees, yet riparian shrubs are key elements of riparian systems and may differ from trees in phenotypic variability and environmental responses. Methods We tested if individuals of a clonal, riparian shrub, Pluchea sericea , collected from provenances spanning a temperature gradient differed in their phenotypes and responses to inundation and to what degree such differences were related to genotype. Plants were subjected to different inundation depths and a subset genotyped. Variables related to growth and resource acquisition were measured and analyzed using hierarchical, multivariate Bayesian linear regressions. Results Individuals from different provenances differed in their phenotypes, but not in their response to inundation. Phenotypes were not related to provenance temperature but were partially governed by genotype. Growth was more strongly influenced by inundation, while resource acquisition was more strongly controlled by genotype. Conclusions Growth and resource acquisition responses in a clonal, riparian shrub are affected by changes to inundation and plant demographics in unique ways. Shrubs appear to differ from trees in their responses to environmental change. Understanding environmental effects on shrubs separately from those of trees will be a key part of evaluating environmental change impacts on riparian ecosystems.

American Journal of Botany

The hydroclimate niche: A tool for predicting and managing riparian plant community responses to streamflow seasonality

Habitat suitability is a consequence of interacting environmental factors. In riparian ecosystems, suitable plant habitat is influenced by interactions between stream hydrology and climate, hereafter referred to as “hydroclimate”. We tested the hypothesis that hydroclimate variables would improve the fit of ecological niche models for a suite of riparian species using occurrence data from the western United States. We focus on the climate conditions (temperature, precipitation and vapor pressure deficit) during the months of lowest and highest streamflow as integrative hydroclimate metrics of resource and stress levels. We found that the inclusion of hydroclimate variables improved model fit for all species in the western USA dataset. We then tested the utility of the improved habitat suitability models by projecting them onto a regulated segment of the Colorado River to assess potential impacts of streamflow seasonality on vegetation metrics of management concern. Species frequency derived from independent survey data in the Colorado River segment was significantly higher for species with predicted suitable habitat than for species without predicted suitable habitat. Under different simulated hydrographs for the Colorado River, overall species richness was predicted to be greatest with peak streamflows during summer, and native-to-non-native species ratios were predicted to be greatest with lowest streamflows in winter. Summer high flows were particularly associated with higher predicted habitat suitability for species that have increased in cover over recent decades (e.g., Pluchea sericea, Baccharis species). We conclude that hydroclimate covariates can be useful tools for predicting how riparian vegetation communities respond to changes in the seasonal timing of low and high streamflows.

River Research and Applications

Functional composition of plant communities mediates biomass effects on ecosystem service recovery across an experimental dryland restoration network

Land degradation can result in a loss of critical ecosystem services that we often seek to restore through re-establishment of desired plant communities. Trait-based approaches have the potential to target specific ecosystem services based on associations between the functional composition of plant communities and ecosystem properties that serve as indicators of those services. The effect of functional composition on ecosystem recovery may depend on the amount of restored plant biomass, itself a supporting service frequently targeted in restoration efforts. Yet, interactions between functional composition and biomass are not formally integrated into trait-based analytical frameworks. We tested the hypothesis that functional composition of plant communities both drives, and interacts with, biomass production to influence indicators of soil functioning and weed suppression across a network of degraded dryland restoration experiments. This networked approach allowed us to identify generalized effects of functional composition on ecosystem recovery across a range of dryland climate conditions. Climate had a substantial effect on ecosystem indicators, with weed cover and soil surface stability increasing in more arid climates, water infiltration increasing with precipitation, and aggregate structure increasing with less freezing. After accounting for climate effects across study sites, we found significant effects of community-weighted mean (CWM) trait values on biomass, particularly a positive effect of leaf carbon-to-nitrogen ratio, and of CWM-biomass interactions on other ecosystem indicators. Cover of exotic species was reduced in restored communities with a combination of low leaf dry matter content and high biomass, soil water infiltration increased with lower specific root length and high biomass, and soil aggregate stability increased with higher root dry matter content and high biomass, among other effects. Functional diversity had no significant effects on any ecosystem indicators. Synthesis : The influence of community functional composition on ecosystem properties increases with community biomass, particularly in disturbed or low productivity systems. This suggests that active management should not only focus on trait values that optimize individual ecosystem indicators but also how those functional strategies are complementary or counter to those that increase biomass.

Functional Ecology

Primary production responses to extreme changes in North American Monsoon precipitation vary by elevation and plant functional composition through time

Primary production in dryland ecosystems is limited by water availability and projected to be strongly affected by future shifts in seasonal precipitation. Warm-season precipitation derived from the North American Monsoon contributes 40% of annual precipitation to dryland ecosystems in the southwestern U.S. and is projected to become more variable. However, there is large uncertainty on whether this variability will be expressed as either extreme wet or dry years and how primary production of different plant functional types will respond across widespread elevation gradients in this region. We experimentally imposed extreme drought and water addition treatments from 2016 – 2020, during which ambient warm-season precipitation declined to reach historic lows, to understand production sensitivity of dominant plant functional types along a 1,000 m elevation gradient. We found that the production responses of plant functional types to monsoon precipitation extremes were dependent on the number of treatment years that occurred across sites along the elevation gradient. C 4 perennial grasses were most responsive to precipitation manipulation treatments, followed by C 3 perennial grasses and annuals, while perennial forbs and shrubs had weak or no responses. C 4 perennial grass reductions due to extreme drought were generally stronger or occurred earlier at low elevation sites, while multi-year extreme drought extended negative effects to C 3 perennial grasses at high elevation, and all sites showed delayed responses to multi-year water addition. We found that the sensitivity of C 3 perennial grass production differed for extreme drought and water addition compared to ambient precipitation at one site, but other sites and plant functional types had similar sensitivities to the different treatment types. Synthesis . The upward advance of primary production responsiveness from single- to multi-year extreme changes in warm-season precipitation suggests more immediate shifts in functional composition and carbon cycling at low elevation, while high elevation ecosystems may become less resistant as the effects of extreme precipitation compound through time.

Arizona