Abstract Background and Aims The success of invasive plants can be attributed to many traits including the ability to adapt to variable environmental conditions. Whether by adaptation, acclimation or phenotypic plasticity, these plants often increase their resource-use efficiency and, consequently, their fitness. The goal of this study was to examine the hydraulic and eco-physiological attributes of sun and shade populations of Pteridium aquilinum, a weedy fern, to determine whether the presence of vessels and other hydraulic attributes affects its success under a variety of light conditions. Methods Hydraulic traits such as cavitation resistance, hydraulic conductivity, photosynthesis and water potential at turgor loss point were measured on fronds from sun and shade populations. Anatomical and structural traits such as conduit diameter and length, stomatal density and vein density were also recorded. Diurnal measures of leaf water potential and stomatal conductance complement these data. Key Results Gas exchange was nearly double in the sun plants, as was water-use efficiency, leaf-specific conductivity, and stomatal and vein density. This was largely achieved by a decrease in leaf area, coupled with higher xylem content. There was no significant difference in petiole cavitation resistance between the sun and shade leaves, nor in xylem-specific conductivity. Hydraulic conduit diameters were nearly equivalent in the two leaf types. Conclusions Shifts in leaf area and xylem content allow P. aquilinum to occupy habitats with full sun, and to adjust its physiology accordingly. High rates of photosynthesis explain in part the success of this fern in disturbed habitats, although no change was observed in intrinsic xylem qualities such as cavitation resistance or conduit length. This suggests that P. aquilinum is constrained by its fundamental body plan, in contrast to seed plants, which show greater capacity for hydraulic adjustment.
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Hydraulic traits that buffer deep‐rooted plants from changes in hydrology and climate
Groundwater‐dependent ecosystems are often defined by the presence of deeply rooted phreatophytic plants. When connected to groundwater, phreatophytes in arid regions decouple ecosystem net primary productivity from precipitation, underscoring a disproportionately high biodiversity and exchange of resources relative to surrounding areas. However, groundwater‐dependent ecosystems are widely threatened due to the effects of water diversions, groundwater abstraction, and higher frequencies of episodic drought and heat waves. The resilience of these ecosystems to shifting ecohydrological–climatological conditions will depend largely on the capacity of dominant, phreatophytic plants to cope with dramatic reductions in water availability and increases in atmospheric water demand. This paper disentangles the broad range of hydraulic traits expressed by phreatophytic vegetation to better understand their capacity to survive or even thrive under shifting ecohydrological conditions. We focus on three elements of plant water relations: (a) hydraulic architecture (including root area to leaf area ratios and rooting depth), (b) xylem structure and function, and (c) stomatal regulation. We place the expression of these traits across a continuum of phreatophytic habits from obligate to semi‐obligate to semi‐facultative to facultative. Although many species occupy multiple phreatophytic niches depending on access to groundwater, we anticipate that populations are largely locally adapted to a narrow range of ecohydrological conditions regardless of gene flow across ecohydrological gradients. Consequently, we hypothesize that reductions in available groundwater and increases in atmospheric water demand will result in either (a) stand replacement of obligate phreatophytic species with more facultative species as a function of widespread mortality in highly groundwater‐dependent populations or (b) directional selection in semi‐obligate and semi‐facultative phreatophytes towards the expression of traits associated with highly facultative phreatophytes in the absence of species replacement. Anticipated shifts in the expression of hydraulic traits may have profound impacts on water cycling processes, species assemblages, and habitat structure of groundwater‐dependent woodlands and riparian forests.
more » « less- NSF-PAR ID:
- 10458173
- Publisher / Repository:
- Wiley Blackwell (John Wiley & Sons)
- Date Published:
- Journal Name:
- Hydrological Processes
- Volume:
- 34
- Issue:
- 2
- ISSN:
- 0885-6087
- Page Range / eLocation ID:
- p. 209-222
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract Drought‐induced groundwater decline and warming associated with climate change are primary threats to dryland riparian woodlands. We used the extreme 2012–2019 drought in southern California as a natural experiment to assess how differences in water‐use strategies and groundwater dependence may influence the drought susceptibility of dryland riparian tree species with overlapping distributions. We analyzed tree‐ring stable carbon and oxygen isotopes collected from two cottonwood species (
Populus trichocarpa andP .fremontii ) along the semi‐arid Santa Clara River. We also modeled tree source water δ18O composition to compare with observed source water δ18O within the floodplain to infer patterns of groundwater reliance. Our results suggest that both species functioned as facultative phreatophytes that used shallow soil moisture when available but ultimately relied on groundwater to maintain physiological function during drought. We also observed apparent species differences in water‐use strategies and groundwater dependence related to their regional distributions.P .fremontii was constrained to more arid river segments and ostensibly used a greater proportion of groundwater to satisfy higher evaporative demand.P .fremontii maintained ∆13C at pre‐drought levels up until the peak of the drought, when trees experienced a precipitous decline in ∆13C. This response pattern suggests that trees prioritized maintaining photosynthetic processes over hydraulic safety, until a critical point. In contrast,P .trichocarpa showed a more gradual and sustained reduction in ∆13C, indicating that drought conditions induced stomatal closure and higher water use efficiency. This strategy may confer drought avoidance forP .trichocarpa while increasing its susceptibility to anticipated climate warming. -
BACKGROUND The availability of nitrogen (N) to plants and microbes has a major influence on the structure and function of ecosystems. Because N is an essential component of plant proteins, low N availability constrains the growth of plants and herbivores. To increase N availability, humans apply large amounts of fertilizer to agricultural systems. Losses from these systems, combined with atmospheric deposition of fossil fuel combustion products, introduce copious quantities of reactive N into ecosystems. The negative consequences of these anthropogenic N inputs—such as ecosystem eutrophication and reductions in terrestrial and aquatic biodiversity—are well documented. Yet although N availability is increasing in many locations, reactive N inputs are not evenly distributed globally. Furthermore, experiments and theory also suggest that global change factors such as elevated atmospheric CO 2 , rising temperatures, and altered precipitation and disturbance regimes can reduce the availability of N to plants and microbes in many terrestrial ecosystems. This can occur through increases in biotic demand for N or reductions in its supply to organisms. Reductions in N availability can be observed via several metrics, including lowered nitrogen concentrations ([N]) and isotope ratios (δ 15 N) in plant tissue, reduced rates of N mineralization, and reduced terrestrial N export to aquatic systems. However, a comprehensive synthesis of N availability metrics, outside of experimental settings and capable of revealing large-scale trends, has not yet been carried out. ADVANCES A growing body of observations confirms that N availability is declining in many nonagricultural ecosystems worldwide. Studies have demonstrated declining wood δ 15 N in forests across the continental US, declining foliar [N] in European forests, declining foliar [N] and δ 15 N in North American grasslands, and declining [N] in pollen from the US and southern Canada. This evidence is consistent with observed global-scale declines in foliar δ 15 N and [N] since 1980. Long-term monitoring of soil-based N availability indicators in unmanipulated systems is rare. However, forest studies in the northeast US have demonstrated decades-long decreases in soil N cycling and N exports to air and water, even in the face of elevated atmospheric N deposition. Collectively, these studies suggest a sustained decline in N availability across a range of terrestrial ecosystems, dating at least as far back as the early 20th century. Elevated atmospheric CO 2 levels are likely a main driver of declines in N availability. Terrestrial plants are now uniformly exposed to ~50% more of this essential resource than they were just 150 years ago, and experimentally exposing plants to elevated CO 2 often reduces foliar [N] as well as plant-available soil N. In addition, globally-rising temperatures may raise soil N supply in some systems but may also increase N losses and lead to lower foliar [N]. Changes in other ecosystem drivers—such as local climate patterns, N deposition rates, and disturbance regimes—individually affect smaller areas but may have important cumulative effects on global N availability. OUTLOOK Given the importance of N to ecosystem functioning, a decline in available N is likely to have far-reaching consequences. Reduced N availability likely constrains the response of plants to elevated CO 2 and the ability of ecosystems to sequester carbon. Because herbivore growth and reproduction scale with protein intake, declining foliar [N] may be contributing to widely reported declines in insect populations and may be negatively affecting the growth of grazing livestock and herbivorous wild mammals. Spatial and temporal patterns in N availability are not yet fully understood, particularly outside of Europe and North America. Developments in remote sensing, accompanied by additional historical reconstructions of N availability from tree rings, herbarium specimens, and sediments, will show how N availability trajectories vary among ecosystems. Such assessment and monitoring efforts need to be complemented by further experimental and theoretical investigations into the causes of declining N availability, its implications for global carbon sequestration, and how its effects propagate through food webs. Responses will need to involve reducing N demand via lowering atmospheric CO 2 concentrations, and/or increasing N supply. Successfully mitigating and adapting to declining N availability will require a broader understanding that this phenomenon is occurring alongside the more widely recognized issue of anthropogenic eutrophication. Intercalibration of isotopic records from leaves, tree rings, and lake sediments suggests that N availability in many terrestrial ecosystems has steadily declined since the beginning of the industrial era. Reductions in N availability may affect many aspects of ecosystem functioning, including carbon sequestration and herbivore nutrition. Shaded areas indicate 80% prediction intervals; marker size is proportional to the number of measurements in each annual mean. Isotope data: (tree ring) K. K. McLauchlan et al. , Sci. Rep. 7 , 7856 (2017); (lake sediment) G. W. Holtgrieve et al. , Science 334 , 1545–1548 (2011); (foliar) J. M. Craine et al. , Nat. Ecol. Evol. 2 , 1735–1744 (2018)more » « less
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Abstract The coordination of traits from individual organs to whole plants is under strong selection because of environmental constraints on resource acquisition and use. However, the tight coordination of traits may provide underlying mechanisms of how locally adapted plant populations can become maladapted because of climate change.
To better understand local adaptation in intraspecific trait coordination, we studied trait variability in the widely distributed foundation tree species,
Populus fremontii using a common garden near the mid‐elevational point of this species distribution. We examined 28 traits encompassing four spectra: phenology, leaf economic spectrum (LES), whole‐tree architecture (Corner's Rule) and wood economic spectrum (WES).Based on adaptive syndrome theory, we hypothesized that trait expression would be coordinated among and within trait spectra, reflecting local adaptation to either exposure to freeze‐thaw conditions in genotypes sourced from high‐elevation populations or exposure to extreme thermal stress in genotypes sourced from low‐elevation populations.
High‐elevation genotypes expressed traits within the phenology and WES that limit frost exposure and tissue damage. Specifically, genotypes sourced from high elevations had later mean budburst, earlier mean budset, higher wood densities, higher bark fractions and smaller xylem vessels than their low‐elevation counterparts. Conversely, genotypes sourced from low elevations expressed traits within the LES that prioritized hydraulic efficiency and canopy thermal regulation to cope with extreme heat exposure, including 40% smaller leaf areas, 67% higher stomatal densities and 34% higher mean theoretical maximum stomatal conductance. Low‐elevation genotypes also expressed a lower stomatal control over leaf water potentials that subsequently dropped to pressures that could induce hydraulic failure.
Synthesis . Our results suggest thatPopulus fremontii expresses a high degree of coordination across multiple trait spectra to adapt to local climate constraints on photosynthetic gas exchange, growth and survival. These results, therefore, increase our mechanistic understanding of local adaptation and the potential effects of climate change that in turn, improves our capacity to identify genotypes that are best suited for future restoration efforts. -
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Abstract As drought variability increases in forests around the globe, it is critical to evaluate and understand ecosystem attributes that ameliorate drought impacts. Trees in arid and semi‐arid ecosystems can sustain tree growth and transpiration during drought by accessing shallow groundwater, yet the extent to which groundwater influences forest growth and transpiration in humid environments has largely been unexplored. We quantified groundwater's influence on tree growth and transpiration in northern humid forests with sandy soils. We hypothesized that even in wet regions, soil droughts occur relatively frequently in forests with sandy soils and result in water stress and reduced tree growth. Further, we hypothesized these reductions in productivity are ameliorated if the forest can access shallow groundwater during dry conditions. We evaluated tree growth responses using tree cores in
Pinus resinosa
