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  1. Abstract

    Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

     
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  2. Abstract

    Understanding the drivers of trait selection is critical for resolving community assembly processes. Here, we test the importance of environmental filtering and trait covariance for structuring the functional traits of understory herbaceous communities distributed along a natural environmental resource gradient that varied in soil moisture, temperature, and nitrogen availability, produced by different topographic positions in the southern Appalachian Mountains.

    To uncover potential differences in community‐level trait responses to the resource gradient, we quantified the averages and variances of both abundance‐weighted and unweighted values for six functional traits (vegetative height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, and leaf δ13C) using 15 individuals of each of the 108 species of understory herbs found at two sites in the southern Appalachians of western North Carolina, USA.

    Environmental variables were better predictors of weighted than unweighted community‐level average trait values for all but height and leaf N, indicating strong environmental filtering of plant abundance. Community‐level variance patterns also showed increased convergence of abundance‐weighted traits as resource limitation became more severe.

    Functional trait covariance patterns based on weighted averages were uniform across the gradient, whereas coordination based on unweighted averages was inconsistent and varied with environmental context. In line with these results, structural equation modeling revealed that unweighted community‐average traits responded directly to local environmental variation, whereas weighted community‐average traits responded indirectly to local environmental variation through trait coordination.

    Our finding that trait coordination is more important for explaining the distribution of weighted than unweighted average trait values along the gradient indicates that environmental filtering acts on multiple traits simultaneously, with abundant species possessing more favorable combinations of traits for maximizing fitness in a given environment.

     
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  3. Abstract

    How much stream temperatures increase within riparian canopy openings and whether stream temperatures cool downstream of these openings both have important policy implications. Past studies of stream cooling downstream of riparian openings have found mixed results including rapid, slow, and no cooling. We collected longitudinal profiles of stream temperatures above, within, and below riparian forest openings along stream segments within otherwise forested riparian conditions to evaluate how sensitivity of stream temperatures to riparian conditions varied across landscape factors. We conducted these temperature surveys across openings in 12 wadeable streams within and near the Upper Little Tennessee River Basin in western North Carolina and northeastern Georgia. Basin areas ranged from 74 to 6,913 ha, and bankfull channel widths varied from 3.4 to 16.4 m. Stream temperatures were collected every 15 min using HOBO® data loggers for 2 weeks in each stream, repeated later in summer in some streams. Reference temperatures were highest in stream reaches at low elevations and with large drainage areas. Stream temperature increases in the middle of riparian gaps were highest when streams drained small high‐elevation watersheds, and increases at the end of openings were highest when the opening length was large relative to watershed size. Downstream from openings, cooling rates were greatest in small, high‐elevation headwater streams and also increased with larger increases in canopy cover. Stream segments that warmed the most within openings also featured higher cooling rates downstream. The data show that stream temperature sensitivity to canopy change is highly dependent on network position and watershed size. A better understanding of stream temperature responses to riparian vegetation may be useful to land managers and landowners prioritizing riparian forest restoration.

     
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  4. Abstract

    Soil organic carbon (SOC) regulates terrestrial ecosystem functioning, provides diverse energy sources for soil microorganisms, governs soil structure, and regulates the availability of organically bound nutrients. Investigators in increasingly diverse disciplines recognize how quantifying SOC attributes can provide insight about ecological states and processes. Today, multiple research networks collect and provide SOC data, and robust, new technologies are available for managing, sharing, and analyzing large data sets. We advocate that the scientific community capitalize on these developments to augment SOC data sets via standardized protocols. We describe why such efforts are important and the breadth of disciplines for which it will be helpful, and outline a tiered approach for standardized sampling of SOC and ancillary variables that ranges from simple to more complex. We target scientists ranging from those with little to no background in soil science to those with more soil‐related expertise, and offer examples of the ways in which the resulting data can be organized, shared, and discoverable.

     
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  5. Abstract

    Dense understory thickets of the native evergreen shrubRhododendron maximumexpanded initially following elimination of American chestnut by the chestnut blight, and later in response to loss of the eastern hemlock due to hemlock woolly adelgid invasion. Rhododendron thickets often blanket streams and their riparian zones, creating cool, low‐light microclimates. To determine the effect of such understory thickets on summer stream temperatures, we removed riparian rhododendron understory on 300 m reaches of two southern Appalachian Mountain headwater streams, while leaving two 300 m reference reaches undisturbed. Overhead canopy was left intact in all four streams, but all streams were selected to have a significant component of dead or dying eastern hemlock in the overstory, creating time‐varying canopy gaps throughout the reach. We continuously monitored temperatures upstream, within and downstream of treatment and reference reaches. Temperatures were monitored in all four streams in the summer before treatments were imposed (2014), and for two summers following treatment (2015, 2016). Temperatures varied significantly across and within streams prior to treatment and across years for the reference streams. After rhododendron removal, increases in summer stream temperatures were observed at some locations within the treatment reaches, but these increases did not persist downstream and varied by watershed, sensor, and year. Significant increases in daily maxima in treatment reaches ranged from 0.9 to 2.6°C. Overhead canopy provided enough shade to prevent rhododendron removal from increasing summer temperatures to levels deleterious to native cold‐water fauna (average summer temperatures remained below 16°C), and local temperature effects were not persistent.

     
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  6. Abstract

    Linking quickflow response to subsurface state can improve our understanding of runoff processes that drive emergent catchment behaviour. We investigated the formation of non‐linear quickflows in three forested headwater catchments and also explored unsaturated and saturated storage dynamics, and likely runoff generation mechanisms that contributed to threshold formation. Our analyses focused on two reference watersheds at the Coweeta Hydrologic Laboratory (CHL) in western North Carolina, USA, and one reference watershed at the Susquehanna Shale Hills Critical Zone Observatory (SHW) in Central Pennsylvania, USA, with available hourly soil moisture, groundwater, streamflow, and precipitation time series over several years. Our study objectives were to characterise (a) non‐linear runoff response as a function of storm characteristics and antecedent conditions, (b) the critical levels of shallow unsaturated and saturated storage that lead to hourly flow response, and (c) runoff mechanisms contributing to rapidly increasing quickflow using measurements of soil moisture and groundwater. We found that maximum hourly rainfall did not significantly contribute to quickflow production in our sites, in contrast to prior studies, due to highly conductive forest soils. Soil moisture and groundwater dynamics measured in hydrologically representative areas of the hillslope showed that variable subsurface states could contribute to non‐linear runoff behaviour. Quickflow generation in watersheds at CHL were dominated by both saturated and unsaturated pathways, but the relative contributions of each pathway varied between catchments. In contrast, quickflow was almost entirely related to groundwater fluctuations at SHW. We showed that co‐located measurements of soil moisture and groundwater supplement threshold analyses providing stronger prediction and understanding of quickflow generation and indicate dominant runoff processes.

     
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  7. Abstract

    Human activities have dramatically altered global patterns of nitrogen (N) and phosphorus (P) availability. This pervasive nutrient pollution is changing basal resource quality in food webs, thereby affecting rates of biological productivity and the pathways of energy and material flow to higher trophic levels.

    Here, we investigate how the stoichiometric quality of basal resources modulates patterns of material flow through food webs by characterizing the effects of experimental N and P enrichment on the trophic basis of macroinvertebrate production and flows of dominant food resources to consumers in five detritus‐based stream food webs.

    After a pre‐treatment year, each stream received N and P at different concentrations for 2 years, resulting in a unique dissolved N:P ratio (target range from 128:1 to 2:1) for each stream. We combined estimates of secondary production and gut contents analysis to calculate rates of material flow from basal resources to macroinvertebrate consumers in all five streams, during all 3 years of study.

    Nutrient enrichment resulted in a 1.5× increase in basal resource flows to primary consumers, with the greatest increases from biofilms and wood. Flows of most basal resources were negatively related to resource C:P, indicating widespread P limitation in these detritus‐based food webs. Nutrient enrichment resulted in a greater proportion of leaf litter, the dominant resource flow‐pathway, being consumed by macroinvertebrates, with that proportion increasing with decreasing leaf litter C:P. However, the increase in efficiency with which basal resources were channelled into metazoan food webs was not propagated to macroinvertebrate predators, as flows of prey did not systematically increase following enrichment and were unrelated to basal resource flows.

    This study suggests that ongoing global increases in N and P supply will increase organic matter flows to metazoan food webs in detritus‐based ecosystems by reducing stoichiometric constraints at basal trophic levels. However, the extent to which those flows are propagated to the highest trophic levels likely depends on responses of individual prey taxa and their relative susceptibility to predation.

     
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  8. Abstract

    Forest canopy water use and carbon cycling traits (WCT) can vary substantially and in spatially organized patterns, with significant impacts on watershed ecohydrology. In many watersheds, WCT may vary systematically along and between hydrologic flowpaths as an adaptation to available soil water, nutrients, and microclimate‐mediated atmospheric water demand. We hypothesize that the emerging patterns of WCT at the hillslope to catchment scale provide a more resistant ecohydrological system, particularly with respect to drought stress, and the maintenance of high levels of productivity. Rather than attempting to address this hypothesis with species‐specific patterns, we outline broader functional WCT groups and explore the sensitivity of water and carbon balances to the representation of canopy WCT functional organization through a modelling approach. We use a well‐studied experimental watershed in North Carolina where detailed mapping of forest community patterns are sufficient to describe WCT functional organization. Ecohydrological models typically use broad‐scale characterizations of forest canopy composition based on remotely sensed information (e.g., evergreen vs. deciduous), which may not adequately represent the range or spatial pattern of functional group WCT at hillslope to watershed scales. We use three different representations of WCT functional organizations: (1) restricting WCT to deciduous/conifer differentiation, (2) utilizing more detailed, but aspatial, information on local forest community composition, and (3) spatially distributed representation of local forest WCT. Accounting for WCT functional organization information improves model performance not only in terms of capturing observed flow regimes (especially watershed‐scale seasonal flow dynamics) but also in terms of representing more detailed canopy ecohydrologic behaviour (e.g., root zone soil moisture, evapotranspiration, and net canopy photosynthesis), especially under dry condition. Results suggest that the well‐known zonation of forest communities over hydrologic gradients is not just a local adaptation but also provides a property that regulates hillslope to catchment‐scale behaviour of water use and drought resistance.

     
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  9. Abstract

    Evaporation of precipitation from plant surfaces, or interception, is a major component of the global water budget. Interception has been measured and/or modelled across a wide variety of forest types; however, most studies have focused on mature, second‐growth forests, and few studies have examined interception processes across forest age classes. We present data on two components of interception, total canopy interception (Ei) and litter interception—that is, Oi + Oehorizon layers—(Eff), across a forest age chronosequence, from 2 years since harvest to old growth. We used precipitation, throughfall, and stemflow collectors to measure total rainfall (P) and estimateEi; and collected litter biomass and modelled litter wetting and drying to estimate evaporative loss from litter. CanopyEi,Pminus throughfall, increased rapidly with forest age and then levelled off to a maximum of 21% ofPin an old‐growth site. Stemflow also varied across stands, with the highest stemflow (~8% ofP) observed in a 12‐year‐old stand with high stem density. ModelledEffwas 4–6% ofPand did not vary across sites. Total stand‐level interception losses (Ei + Eff) were best predicted by stand age (R2 = 0.77) rather than structural parameters such as basal area (R2 = 0.49) or leaf area (R2 < 0.01). Forest age appears to be an important driver of interception losses from forested mountain watersheds even when stand‐level structural variables are similar. These results will contribute to our understanding of water budgets across the broader matrix of forest ages that characterize the modern forest landscape.

     
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  10. Abstract

    Catchments with minimal disturbance usually have low dissolved inorganic nitrogen (DIN) export, but disturbances and anthropogenic inputs result in elevated DIN concentration and export and eutrophication of downstream ecosystems. We studied streams in the southern Appalachian Mountains, USA, an area dominated by hardwood deciduous forest but with areas of valley agriculture and increasing residential development. We collected weekly grab samples and storm samples from nine small catchments and three river sites. Most discharge occurred at baseflow, with baseflow indices ranging from 69% to 95%. We identified three seasonal patterns of baseflow DIN concentration. Streams in mostly forested catchments had low DIN with bimodal peaks, and summer peaks were greater than winter peaks. Streams with more agriculture and development also had bimodal peaks; however, winter peaks were the highest. In streams draining catchments with more residential development, DIN concentration had a single peak, greatest in winter and lowest in summer. Three methods for estimating DIN export produced consistent results. Annual DIN export ranged from less than 200 g ha−1 year−1for the less disturbed catchments to over 2,000 g ha−1 year−1in the catchments with the least forest area. Land cover was a strong predictor of DIN concentration but less significant for predicting DIN export. The two forested reference catchments appeared supply limited, the most residential catchment appeared transport limited, and export for the other catchments was significantly related to discharge. In all streams, baseflow DIN export exceeded stormflow export. Morphological and climatological variation among watersheds created complexities unexplainable by land cover. Nevertheless, regression models developed using land cover data from the small catchments reasonably predicted concentration and export for receiving rivers. Our results illustrate the complexity of mechanisms involved in DIN export in a region with a mosaic of climate, geology, topography, soils, vegetation, and past and present land use.

     
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