Spatial, temporal, and biological factors influencing plant responses to deicing salt in roadside bioinfiltration basins.

Plants are arguably the most visible components of stormwater bioretention basins and play key roles in stabilizing soils and removing water through transpiration. In regions with cold winters, bioretention basins along roadways can receive considerable quantities of deicing salt, much of which migrates out of the systems prior to the onset of plant growth but the rest remains in the soil. The resulting effects on plants presumably vary with time (due to annual weather patterns), space (because stormwater exposure is location-dependent), and biology (because plant taxa differ in their salt tolerance). The goal of this study was to investigate the magnitude of deicing salt's effects on bioretention plants and how it varies with spatial, temporal, and biological factors. The study took place in a set of five bioretention basins in Philadelphia, USA that receive runoff from a major highway. Over a five-year period, the electrical conductivity (EC) of influent stormwater frequently exceeded 1 mS cm-1 in winter, and occasionally surpassed that of seawater (∼50 mS cm-1). In both of the years when soil EC was measured as well, it remained elevated through all spring months, especially near basin inlets and centers. Mortality of nine plant taxa ranged widely after three years (0-90%), with rankings largely corresponding to salt tolerances. Moreover, leaf areas and/or crown volumes were strongly reduced in proportion to stormwater exposure in seven of these taxa. In the three taxa evaluated for tissue concentrations of 14 potentially toxic elements (Hemerocallis 'Happy Returns', Iris 'Caesar's Brother', and Cornus sericea 'Cardinal'), only sodium consistently exceeded the toxicity limit for salt intolerant plants (500 mg kg-1). However, exceedance of the sodium toxicity limit was associated with plants' topographic positions, with median concentrations greatest in the bottom of basins and least on basin rims. This study demonstrates that deicing salts can have detrimental effects on plants in bioretention basins, with the strongest effects likely to occur in years with the greatest snowfall (and therefore deicing salt use), in portions of basins with greatest stormwater exposure (typically around inlets and centers), and in plants with minimal salinity tolerance. Our results therefore underscore the value of installing salt-tolerant taxa in basins likely to experience any frequency of deicing salt exposure.

Canopy composition drives variability in urban growing season length more than the heat island effect.

The elevated heat of urban areas compared to their surroundings makes humid temperate cities a useful preview of future climate effects on natural forest phenology. The utility of this proxy rests on the expectation that trees in urban areas alter their phenology in response to warmer site conditions in spring and fall. However, it is possible that apparent lengthening of the growing season is instead governed by human-driven tree species selection and plant functional type (PFT; trees, shrubs, turfgrass) heterogeneity typical of managed landscapes. Without the use of highly spatially and temporally resolved remote sensing data, the roles of tree taxonomy and local site characteristics (e.g., impervious cover) in controlling phenology remain confounded. To understand the drivers of earlier start of season (SOS) and later end of season (EOS) among urban trees, we estimated individual tree phenology using >130 high-resolution satellite images per year (2018-2020) for ~10,000 species-labeled trees in Washington, DC. We found that species identity alone accounted for 4× more variability in the timing of SOS and EOS compared with a tree's planting location characteristics. Additionally, the urban mix of PFTs may be more responsible for apparent advances in SOS (by between 1.8 ± 1.3 and 3.5 ± 1.3 days) than heat per se. The results of this study caution against associating longer growing seasons in cities-observed in moderate to coarse resolution remote sensing imagery-to within-species phenological plasticity and demonstrate the power of high-resolution satellite data for tracking tree phenology in biodiverse environments.

Systematic review of quantitative preference studies of treatments for rheumatoid arthritis among patients and at-risk populations.

Treatments used for rheumatoid arthritis (RA) are under investigation for their efficacy to prevent RA in at risk groups. It is therefore important to understand treatment preferences of those at risk. We systematically reviewed quantitative preference studies of drugs to treat, or prevent RA, to inform the design of further studies and trials of RA prevention. Stated preference studies for RA treatment or prevention were identified through a search of five databases. Study characteristics and results were extracted, and the relative importance of different types of treatment attributes was compared across populations. Twenty three studies were included 20 of RA treatments (18 of patients; 2 of the general public) and 3 prevention studies with first-degree relatives (FDRs). Benefits, risks, administration method and cost (when included) were important determinants of treatment choice. A benefit was more important than a risk attribute in half of the studies of RA treatment that included a benefit attribute and 2/3 studies of RA prevention. There was variability in the relative importance of attributes across the few prevention studies. In studies with non-patient participants, attributes describing confidence in treatment effectiveness/safety were more important determinants of choice than in studies with patients. Most preference studies relating to RA are of treatments for established RA. Few studies examine preferences for treatments to prevent RA. Given intense research focus on RA prevention, additional preference studies in this context are needed. Variation in treatment preferences across different populations is not well understood and direct comparisons are needed.

Biological invasions and climate change amplify each other's effects on dryland degradation.

Climate models predict that, in the coming decades, many arid regions will experience increasingly hot conditions and will be affected more frequently by drought. These regions are also experiencing rapid vegetation change, notably invasion by exotic grasses. Invasive grasses spread rapidly into native desert ecosystems due, in particular, to interannual variability in precipitation and periodic fires. The resultant destruction of non-fire-adapted native shrub and grass communities and of the inherent soil resource heterogeneity can yield invader-dominated grasslands. Moreover, recurrent droughts are expected to cause widespread physiological stress and mortality of both invasive and native plants, as well as the loss of soil resources. However, the magnitude of these effects may differ between invasive and native grasses, especially under warmer conditions, rendering the trajectory of vegetated communities uncertain. Using the Biosphere 2 facility in the Sonoran Desert, we evaluated the viability of these hypothesized relationships by simulating combinations of drought and elevated temperature (+5°C) and assessing the ecophysiological and mortality responses of both a dominant invasive grass (Pennisetum ciliare or buffelgrass) and a dominant native grass (Heteropogan contortus or tanglehead). While both grasses survived protracted drought at ambient temperatures by inducing dormancy, drought under warmed conditions exceeded the tolerance limits of the native species, resulting in greater and more rapid mortality than exhibited by the invasive. Thus, two major drivers of global environmental change, biological invasion and climate change, can be expected to synergistically accelerate ecosystem degradation unless large-scale interventions are enacted.

Responses of stomatal features and photosynthesis to porewater N enrichment and elevated atmospheric CO2 in Phragmites australis, the common reed.

PREMISE: Biological invasions increasingly threaten native biodiversity and ecosystem services. One notable example is the common reed, Phragmites australis, which aggressively invades North American salt marshes. Elevated atmospheric CO2 and nitrogen pollution enhance its growth and facilitate invasion because P. australis responds more strongly to these enrichments than do native species. We investigated how modifications to stomatal features contribute to strong photosynthetic responses to CO2 and nitrogen enrichment in P. australis by evaluating stomatal shifts under experimental conditions and relating them to maximal stomatal conductance (gwmax ) and photosynthetic rates. METHODS: Plants were grown in situ in open-top chambers under ambient and elevated atmospheric CO2 (eCO2 ) and porewater nitrogen (Nenr ) in a Chesapeake Bay tidal marsh. We measured light-saturated carbon assimilation rates (Asat ) and stomatal characteristics, from which we calculated gwmax and determined whether CO2 and Nenr altered the relationship between gwmax and Asat . RESULTS: eCO2 and Nenr enhanced both gwmax and Asat , but to differing degrees; gwmax was more strongly influenced by Nenr through increases in stomatal density while Asat was more strongly stimulated by eCO2 . There was a positive relationship between gwmax and Asat that was not modified by eCO2 or Nenr , individually or in combination. CONCLUSIONS: Changes in stomatal features co-occur with previously described responses of P. australis to eCO2 and Nenr . Complementary responses of stomatal length and density to these global change factors may facilitate greater stomatal conductance and carbon gain, contributing to the invasiveness of the introduced lineage.

Rapid recovery of carbon cycle processes after the cessation of chronic nutrient enrichment.

Salt marshes provide critical ecosystem services including some of the highest rates of carbon storage on Earth. However, many salt marshes receive very high nutrient loads and there is a growing body of evidence indicating that this nutrient enrichment alters carbon cycle processes. While many restoration plans prioritize nutrient management in their efforts to conserve salt marsh ecosystems, there has been little empirical investigation of the capacity for carbon cycle processes to recover once nutrient loading is reduced. To address this, we compared rates of greenhouse gas fluxes (i.e., CO2 and methane) measured using static chambers, and soil organic matter decomposition, using both litter bags and the Tea Bag Index (TBI), during the last two years of a long-term, ecosystem-scale nutrient enrichment experiment (2015-2016) as well as in the first two years of recovery post-enrichment (2017-2018). We found that both ecosystem respiration (Reco) and decomposition processes (i.e., rhizome decomposition and soil organic matter stabilization) were enhanced by nutrient enrichment, but returned to reference ecosystem levels within the first year following the cessation of nutrient enrichment and remained at reference levels in the second year. These results suggest that management practices intended to reduce nutrient loads in coastal systems may, in fact, allow for rapid recovery of carbon cycle processes, potentially restoring the high carbon sequestration rates of these blue carbon ecosystems.

Decadal-scale shifts in soil hydraulic properties as induced by altered precipitation.

Soil hydraulic properties influence the partitioning of rainfall into infiltration versus runoff, determine plant-available water, and constrain evapotranspiration. Although rapid changes in soil hydraulic properties from direct human disturbance are well documented, climate change may also induce such shifts on decadal time scales. Using soils from a 25-year precipitation manipulation experiment, we found that a 35% increase in water inputs substantially reduced infiltration rates and modestly increased water retention. We posit that these shifts were catalyzed by greater pore blockage by plant roots and reduced shrink-swell cycles. Given that precipitation regimes are expected to change at accelerating rates globally, shifts in soil structure could occur over broad regions more rapidly than expected and thus alter water storage and movement in numerous terrestrial ecosystems.

Fine-root traits are linked to species dynamics in a successional plant community.

Despite the importance of fine roots for the acquisition of soil resources such as nitrogen and water, the study of linkages between traits and both population and community dynamics remains focused on aboveground traits. We address this gap by investigating associations between belowground traits and metrics of species dynamics. Our analysis included 85 species from a long-term data set on the transition from old field to forest in eastern North America (the Buell-Small Succession Study) and the new Fine-Root Ecology Database. Given the prominent roles of life form (woody vs. non-woody) and species origin (native vs. exotic) in defining functional relationships, we also assessed whether traits or their relationships with species dynamics differed for these groups. Species that reached their peak abundance early in succession had fine-root traits corresponding to resource acquisitive strategies (i.e., they were thinner, less dense, and had higher nitrogen concentrations) while species that peaked progressively later had increasingly conservative strategies. In addition to having more acquisitive root traits than native species, exotics diverged from the above successional trend, having consistently thinner fine roots regardless of the community context. Species with more acquisitive fine-root morphologies typically had faster rates of abundance increase and achieved their maximal rates in fewer years. Decreasing soil nutrient availability and increasing belowground competition may become increasingly strong filters in successional communities, acting on root traits to promote a transition from acquisitive to conservative foraging. However, disturbances that increase light and soil resource availability at local scales may allow acquisitive species, especially invasive exotics, to continue colonizing late into the community transition to forest.

Maintenance of photosynthesis by Betula populifolia in metal contaminated soils.

Improving our understanding of plant responses to elevated trace metal concentrations under field conditions will enhance restoration and urban greening practices in settings with contaminated soils. This study examined the effects of trace metal pollution on the leaf gas exchange rates of mature, field-grown Betula populifolia Marsh. (gray birch) trees, additionally assessing whether elevated temperature and drought compounded the effects of trace metal contamination. The study compared B. populifolia growing in areas of comparatively high and low trace metal loads (HML and LML, respectively) within a former rail yard at Liberty State Park in Jersey City, New Jersey, USA. Gas exchange parameters were determined monthly from May through September in 2014 and 2015 using a portable photosynthesis system. The effects of drought and high temperature were assessed during a short heat wave in July 2015 and via a manipulative experiment, respectively. During a few of the measurement months, some parameters differed significantly between the LML and HML groups. However, when considered over the entire study period, no significant differences in biophysical parameters were observed between groups. The photosynthetic capacity of B. populifolia thus appears to be fairly robust across this site's steep gradient of trace metal contamination. Nonetheless, leaf mass per unit area was significantly lower in the HML group, indicating that metal loads affected resource allocation within trees. Also, immediately following the heat wave in 2015, intrinsic water use efficiency declined significantly in the HML group, suggesting that extreme climatic conditions can have a disproportionate effect on the physiological performance of plants growing in metal contaminated soils.

Nitrogen uptake kinetics and saltmarsh plant responses to global change.

Coastal wetlands are important carbon sinks globally, but their ability to store carbon hinges on their nitrogen (N) supply and N uptake dynamics of dominant plant species. In terrestrial ecosystems, uptake of nitrate (NO3-) and ammonium (NH4+) through roots can strongly influence N acquisition rates and their responses to environmental factors such as rising atmospheric CO2 and eutrophication. We examined the 15N uptake kinetics of three dominant plant species in North American coastal wetlands (Spartina patens, C4 grass; Phragmites australis, C3 grass; Schoenoplectus americanus, C3 sedge) under ambient and elevated CO2 conditions. We further related our results to the productivity response of these species in two long-term field experiments. S. patens had the greatest uptake rates for NO3- and NH4+ under ambient conditions, suggesting that N uptake kinetics may underlie its strong productivity response to N in the field. Elevated CO2 increased NH4+ and NO3- uptake rates for S. patens, but had negative effects on NO3- uptake rates in P. australis and no effects on S. americanus. We suggest that N uptake kinetics may explain differences in plant community composition in coastal wetlands and that CO2-induced shifts, in combination with N proliferation, could alter ecosystem-scale productivity patterns of saltmarshes globally.

Cosmopolitan Species As Models for Ecophysiological Responses to Global Change: The Common Reed Phragmites australis.

Phragmites australis is a cosmopolitan grass and often the dominant species in the ecosystems it inhabits. Due to high intraspecific diversity and phenotypic plasticity, P. australis has an extensive ecological amplitude and a great capacity to acclimate to adverse environmental conditions; it can therefore offer valuable insights into plant responses to global change. Here we review the ecology and ecophysiology of prominent P. australis lineages and their responses to multiple forms of global change. Key findings of our review are that: (1) P. australis lineages are well-adapted to regions of their phylogeographic origin and therefore respond differently to changes in climatic conditions such as temperature or atmospheric CO2; (2) each lineage consists of populations that may occur in geographically different habitats and contain multiple genotypes; (3) the phenotypic plasticity of functional and fitness-related traits of a genotype determine the responses to global change factors; (4) genotypes with high plasticity to environmental drivers may acclimate or even vastly expand their ranges, genotypes of medium plasticity must acclimate or experience range-shifts, and those with low plasticity may face local extinction; (5) responses to ancillary types of global change, like shifting levels of soil salinity, flooding, and drought, are not consistent within lineages and depend on adaptation of individual genotypes. These patterns suggest that the diverse lineages of P. australis will undergo intense selective pressure in the face of global change such that the distributions and interactions of co-occurring lineages, as well as those of genotypes within-lineages, are very likely to be altered. We propose that the strong latitudinal clines within and between P. australis lineages can be a useful tool for predicting plant responses to climate change in general and present a conceptual framework for using P. australis lineages to predict plant responses to global change and its consequences.

Nutrient foraging strategies are associated with productivity and population growth in forest shrubs.

Background and Aims: Temperate deciduous forest understoreys are experiencing widespread changes in community composition, concurrent with increases in rates of nitrogen supply. These shifts in plant abundance may be driven by interspecific differences in nutrient foraging (i.e. conservative vs. acquisitive strategies) and, thus, adaptation to contemporary nutrient loading conditions. This study sought to determine if interspecific differences in nutrient foraging could help explain patterns of shrub success and decline in eastern North American forests. Methods: Using plants grown in a common garden, fine root traits associated with nutrient foraging were measured for six shrub species. Traits included the mean and skewness of the root diameter distribution, specific root length (SRL), C:N ratio, root tissue density, arbuscular mycorrhizal colonization and foraging precision. Above- and below-ground productivity were also determined for the same plants, and population growth rates were estimated using data from a long-term study of community dynamics. Root traits were compared among species and associations among root traits, measures of productivity and rates of population growth were evaluated. Key Results: Species fell into groups having thick or thin root forms, which correspond to conservative vs. acquisitive nutrient foraging strategies. Interspecific variation in root morphology and tissue construction correlated with measures of productivity and rates of cover expansion. Of the four species with acquisitive traits, three were introduced species that have become invasive in recent decades, and the fourth was a weedy native. In contrast, the two species with conservative traits were historically dominant shrubs that have declined in abundance in eastern North American forests. Conclusions: In forest understoreys of eastern North America, elevated nutrient availability may impose a filter on species success in addition to above-ground processes such as herbivory and overstorey canopy conditions. Shrubs that have root traits associated with rapid uptake of soil nutrients may be more likely to increase in abundance, while species without such traits may be less likely to keep pace with more productive species.

Nitrogen-mediated effects of elevated CO2 on intra-aggregate soil pore structure.

Soil pore structure has a strong influence on water retention, and is itself influenced by plant and microbial dynamics such as root proliferation and microbial exudation. Although increased nitrogen (N) availability and elevated atmospheric CO2 concentrations (eCO2 ) often have interacting effects on root and microbial dynamics, it is unclear whether these biotic effects can translate into altered soil pore structure and water retention. This study was based on a long-term experiment (7 yr at the time of sampling) in which a C4 pasture grass (Paspalum notatum) was grown on a sandy loam soil while provided factorial additions of N and CO2 . Through an analysis of soil aggregate fractal properties supported by 3D microtomographic imagery, we found that N fertilization induced an increase in intra-aggregate porosity and a simultaneous shift toward greater accumulation of pore space in larger aggregates. These effects were enhanced by eCO2 and yielded an increase in water retention at pressure potentials near the wilting point of plants. However, eCO2 alone induced changes in the opposite direction, with larger aggregates containing less pore space than under control conditions, and water retention decreasing accordingly. Results on biotic factors further suggested that organic matter gains or losses induced the observed structural changes. Based on our results, we postulate that the pore structure of many mineral soils could undergo N-dependent changes as atmospheric CO2 concentrations rise, having global-scale implications for water balance, carbon storage, and related rhizosphere functions.

Box-Counting Dimension Revisited: Presenting an Efficient Method of Minimizing Quantization Error and an Assessment of the Self-Similarity of Structural Root Systems.

Fractal dimension (FD), estimated by box-counting, is a metric used to characterize plant anatomical complexity or space-filling characteristic for a variety of purposes. The vast majority of published studies fail to evaluate the assumption of statistical self-similarity, which underpins the validity of the procedure. The box-counting procedure is also subject to error arising from arbitrary grid placement, known as quantization error (QE), which is strictly positive and varies as a function of scale, making it problematic for the procedure's slope estimation step. Previous studies either ignore QE or employ inefficient brute-force grid translations to reduce it. The goals of this study were to characterize the effect of QE due to translation and rotation on FD estimates, to provide an efficient method of reducing QE, and to evaluate the assumption of statistical self-similarity of coarse root datasets typical of those used in recent trait studies. Coarse root systems of 36 shrubs were digitized in 3D and subjected to box-counts. A pattern search algorithm was used to minimize QE by optimizing grid placement and its efficiency was compared to the brute force method. The degree of statistical self-similarity was evaluated using linear regression residuals and local slope estimates. QE, due to both grid position and orientation, was a significant source of error in FD estimates, but pattern search provided an efficient means of minimizing it. Pattern search had higher initial computational cost but converged on lower error values more efficiently than the commonly employed brute force method. Our representations of coarse root system digitizations did not exhibit details over a sufficient range of scales to be considered statistically self-similar and informatively approximated as fractals, suggesting a lack of sufficient ramification of the coarse root systems for reiteration to be thought of as a dominant force in their development. FD estimates did not characterize the scaling of our digitizations well: the scaling exponent was a function of scale. Our findings serve as a caution against applying FD under the assumption of statistical self-similarity without rigorously evaluating it first.

Belowground advantages in construction cost facilitate a cryptic plant invasion.

The energetic cost of plant organ construction is a functional trait that is useful for understanding carbon investment during growth (e.g. the resource acquisition vs. tissue longevity tradeoff), as well as in response to global change factors like elevated CO2 and N. Despite the enormous importance of roots and rhizomes in acquiring soil resources and responding to global change, construction costs have been studied almost exclusively in leaves. We sought to determine how construction costs of aboveground and belowground organs differed between native and introduced lineages of a geographically widely dispersed wetland plant species (Phragmites australis) under varying levels of CO2 and N. We grew plants under ambient and elevated atmospheric CO2, as well as under two levels of soil nitrogen. We determined construction costs for leaves, stems, rhizomes and roots, as well as for whole plants. Across all treatment conditions, the introduced lineage of Phragmites had a 4.3 % lower mean rhizome construction cost than the native. Whole-plant construction costs were also smaller for the introduced lineage, with the largest difference in sample means (3.3 %) occurring under ambient conditions. In having lower rhizome and plant-scale construction costs, the introduced lineage can recoup its investment in tissue construction more quickly, enabling it to generate additional biomass with the same energetic investment. Our results suggest that introduced Phragmites has had an advantageous tissue investment strategy under historic CO2 and N levels, which has facilitated key rhizome processes, such as clonal spread. We recommend that construction costs for multiple organ types be included in future studies of plant carbon economy, especially those investigating global change.

Descendant root volume varies as a function of root type: estimation of root biomass lost during uprooting in Pinus pinaster.

Root systems of woody plants generally display a strong relationship between the cross-sectional area or cross-sectional diameter (CSD) of a root and the dry weight of biomass (DWd) or root volume (Vd) that has grown (i.e., is descendent) from a point. Specification of this relationship allows one to quantify root architectural patterns and estimate the amount of material lost when root systems are extracted from the soil. However, specifications of this relationship generally do not account for the fact that root systems are comprised of multiple types of roots. We assessed whether the relationship between CSD and Vd varies as a function of root type. Additionally, we sought to identify a more accurate and time-efficient method for estimating missing root volume than is currently available. We used a database that described the 3D root architecture of Pinus pinaster root systems (5, 12, or 19 years) from a stand in southwest France. We determined the relationship between CSD and Vd for 10,000 root segments from intact root branches. Models were specified that did and did not account for root type. The relationships were then applied to the diameters of 11,000 broken root ends to estimate the volume of missing roots. CSD was nearly linearly related to the square root of Vd, but the slope of the curve varied greatly as a function of root type. Sinkers and deep roots tapered rapidly, as they were limited by available soil depth. Distal shallow roots tapered gradually, as they were less limited spatially. We estimated that younger trees lost an average of 17% of root volume when excavated, while older trees lost 4%. Missing volumes were smallest in the central parts of root systems and largest in distal shallow roots. The slopes of the curves for each root type are synthetic parameters that account for differentiation due to genetics, soil properties, or mechanical stimuli. Accounting for this differentiation is critical to estimating root loss accurately.

Jatropha curcas L. root structure and growth in diverse soils.

Unlike most biofuel species, Jatropha curcas has promise for use in marginal lands, but it may serve an additional role by stabilizing soils. We evaluated the growth and structural responsiveness of young J. curcas plants to diverse soil conditions. Soils included a sand, a sandy-loam, and a clay-loam from eastern Mexico. Growth and structural parameters were analyzed for shoots and roots, although the focus was the plasticity of the primary root system architecture (the taproot and four lateral roots). The sandy soil reduced the growth of both shoot and root systems significantly more than sandy-loam or clay-loam soils; there was particularly high plasticity in root and shoot thickness, as well as shoot length. However, the architecture of the primary root system did not vary with soil type; the departure of the primary root system from an index of perfect symmetry was 14 ± 5% (mean ± standard deviation). Although J. curcas developed more extensively in the sandy-loam and clay-loam soils than in sandy soil, it maintained a consistent root to shoot ratio and root system architecture across all types of soil. This strong genetic determination would make the species useful for soil stabilization purposes, even while being cultivated primarily for seed oil.

Functional morphology underlies performance differences among invasive and non-invasive ruderal Rubus species.

The ability of some introduced plant species to outperform native species under altered resource conditions makes them highly productive in ecosystems with surplus resources. However, ruderal native species are also productive when resources are available. The differences in abundance among invasive and non-invasive ruderal plants may be related to differences in ability to maintain access to or store resources for continual use. For a group of ruderal species in the Pacific Northwest of North America (invasive Rubus armeniacus; non-invasive R. ursinus, R. parviflorus, R. spectabilis, and Rosa nutkana), we sought to determine whether differences in functional morphological traits, especially metrics of water access and storage, were consistent with differences in water conductance and growth rate. We also investigated the changes in these traits in response to abundant vs. limited water availability. Rubus armeniacus had among the largest root systems and cane cross-sectional areas, the lowest cane tissue densities, and the most plastic ratios of leaf area to plant mass and of xylem area to leaf area, often sharing its rank with R. ursinus or Rosa nutkana. These three species had the highest water conductance and relative growth rates, though Rubus armeniacus grew the most rapidly when water was not limited. Our results suggest that water access and storage abilities vary with morphology among the ruderal species investigated, and that these abilities, in combination, are greatest in the invasive. In turn, functional morphological traits allow R. armeniacus to maintain rapid gas exchange rates during the dry summers in its invaded range, conferring on it high productivity.