Joshua S. Caplan

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.

Global change accelerates carbon assimilation by a wetland ecosystem engineer

The primary productivity of coastal wetlands is changing dramatically in response to rising atmospheric carbon dioxide (CO2) concentrations, nitrogen (N) enrichment, and invasions by novel species, potentially altering their ecosystem services and resilience to sea level rise. In order to determine how these interacting global change factors will affect coastal wetland productivity, we quantified growing-season carbon assimilation (≈gross primary productivity, or GPP) and carbon retained in living plant biomass (≈net primary productivity, or NPP) of North American mid-Atlantic saltmarshes invaded by Phragmites australis (common reed) under four treatment conditions: two levels of CO2 (ambient and +300 ppm) crossed with two levels of N (0 and 25 g N added m−2 yr−1). For GPP, we combined descriptions of canopy structure and leaf-level photosynthesis in a simulation model, using empirical data from an open-top chamber field study. Under ambient CO2 and low N loading (i.e., the Control), we determined GPP to be 1.66 ± 0.05 kg C m−2 yr−1 at a typical Phragmites stand density. Individually, elevated CO2 and N enrichment increased GPP by 44 and 60%, respectively. Changes under N enrichment came largely from stimulation to carbon assimilation early and late in the growing season, while changes from CO2 came from stimulation during the early and mid-growing season. In combination, elevated CO2 and N enrichment increased GPP by 95% over the Control, yielding 3.24 ± 0.08 kg C m−2 yr−1. We used biomass data to calculate NPP, and determined that it represented 44%–60% of GPP, with global change conditions decreasing carbon retention compared to the Control. Our results indicate that Phragmites invasions in eutrophied saltmarshes are driven, in part, by extended phenology yielding 3.1× greater NPP than native marsh. Further, we can expect elevated CO2 to amplify Phragmites productivity throughout the growing season, with potential implications including accelerated spread and greater carbon storage belowground.

Belowground advantages in construction cost facilitate a cryptic plant invasion

Energetic costs of tissue construction were compared in two subspecies of Phragmites australis, the common reed – namely the primary native and introduced lineages in North America. Caplan et al. report that the introduced lineage has lower construction costs than the native under all environmental conditions assessed, driven mainly by its lower cost rhizomes. These results highlight the fact that belowground energetics, which are seldom investigated, can influence the performance advantages that drive many plant invasions. The authors also demonstrate that tissue construction costs in organs not typically assessed can shift with global change, suggesting that they may have increasingly important implications into the future.

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.

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.

Influence of variable precipitation on coastal water quality in southern California.

OBJECTIVES To examine the consequences of changing precipitation levels on southern Californias recreational coastal water quality, and compare the responses of watersheds with differing levels of urban development. METHODS The geo-temporal relationship for six years (2000-2005) of precipitation levels, discharge rates for the ten primary waterways, and coastal water bacteria concentrations at seventy-eight southern California beaches were examined. RESULTS Precipitation levels, river-creek discharge rates, and coastal water bacteria concentrations were significantly correlated (p < 0.01) for all ten watersheds investigated. Water bacteria concentrations significantly increased with higher levels of precipitation across 95% of the seventy-eight beaches investigated. A heavily developed watershed had significantly higher median bacteria concentrations (186 cfu) in the adjoining coastal waters compared to an undeveloped watershed (10 cfu) of similar size. CONCLUSIONS Precipitation and ensuing runoff strongly control the rate of polluted water delivered to most beaches in southern California. Variable precipitation generates a greater response in coastal water bacteria concentrations in developed watersheds compared to undeveloped areas. Projected declines in regional precipitation as a consequence of climate change may result in less contaminated water delivered to coastal waters, thus decreasing risk of water associated illnesses during winter months.

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.

Allometry data and equations for coastal marsh plants

Coastal marshes are highly valued for ecosystem services such as protecting inland habitats from storms, sequestering carbon, removing nutrients and other pollutants from surface water, and providing habitat for fish, shellfish, and birds. Because plants largely determine the structure and function of coastal marshes, quantifying plant biomass is essential for evaluating these ecosystem services, understanding the biogeochemical processes that regulate ecosystem function, and forecasting tidal wetland responses to accelerated sea level rise. Allometry is a convenient and efficient technique for nondestructive estimation of plant biomass, and it is commonly used in studies of carbon and nitrogen cycles, energy flows, and marsh surface elevation change. We present plant allometry data and models developed for three long-term experiments at the Smithsonian Global Change Research Wetland, a brackish marsh in the Rhode River subestuary of the Chesapeake Bay. The dataset contains 9,771 measurements of stem height, dry mass, and (in 9638 cases) stem width across 11 plant species. The vast majority of observations are for Schoenoplectus americanus (8430) and Phragmites australis (311), with fewer observations for other common species: Amaranthus cannabinus, Atriplex patula, Iva frutescens, Kosteletzkya virginica, Polygonum hydropiper, Solidago sempervirens, Spartina alterniflora, Spartina cynosuroides, and Typha angustifolia. Allometric relationships take the form of linear regressions of biomass (transformed using the Box-Cox procedure) on either stem height and width, or on stem height alone. Allometric relationships for Schoenoplectus americanus were not meaningfully altered by elevated CO2 , N enrichment, the community context, interannual variation in climate, or year, showing that a single equation can be used across a broad range of conditions for this species. Archived files include: (1) raw data used to derive allometric equations for each species, (2) reports and evaluations of the allometric equations we derived from the data, and (3) R code with which our derivations can be replicated. Methodological details of our experiments, data collection efforts, and statistical modeling are described in the metadata. The allometric equations can be used for biomass estimation in empirical and modeling studies of North American coastal wetlands, and the data can be used in ecological studies of terrestrial plant allometry.

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.

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 procedures 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.