A research agenda for nonvascular photoautotrophs under climate change.

Nonvascular photoautotrophs (NVP), including bryophytes, lichens, terrestrial algae, and cyanobacteria, are increasingly recognized as being essential to ecosystem functioning in many regions of the world. Current research suggests that climate change may pose a substantial threat to NVP, but the extent to which this will affect the associated ecosystem functions and services is highly uncertain. Here, we propose a research agenda to address this urgent question, focusing on physiological and ecological processes that link NVP to ecosystem functions while also taking into account the substantial taxonomic diversity across multiple ecosystem types. Accordingly, we developed a new categorization scheme, based on microclimatic gradients, which simplifies the high physiological and morphological diversity of NVP and world-wide distribution with respect to several broad habitat types. We found that habitat-specific ecosystem functions of NVP will likely be substantially affected by climate change, and more quantitative process understanding is required on: (1) potential for acclimation; (2) response to elevated CO2 ; (3) role of the microbiome; and (4) feedback to (micro)climate. We suggest an integrative approach of innovative, multimethod laboratory and field experiments and ecophysiological modelling, for which sustained scientific collaboration on NVP research will be essential.

Shower thoughts: why scientists should spend more time in the rain.

Stormwater is a vital resource and dynamic driver of terrestrial ecosystem processes. However, processes controlling interactions during and shortly after storms are often poorly seen and poorly sensed when direct observations are substituted with technological ones. We discuss how human observations complement technological ones and the benefits of scientists spending more time in the storm. Human observation can reveal ephemeral storm-related phenomena such as biogeochemical hot moments, organismal responses, and sedimentary processes that can then be explored in greater resolution using sensors and virtual experiments. Storm-related phenomena trigger lasting, oversized impacts on hydrologic and biogeochemical processes, organismal traits or functions, and ecosystem services at all scales. We provide examples of phenomena in forests, across disciplines and scales, that have been overlooked in past research to inspire mindful, holistic observation of ecosystems during storms. We conclude that technological observations alone are insufficient to trace the process complexity and unpredictability of fleeting biogeochemical or ecological events without the shower thoughts produced by scientists' human sensory and cognitive systems during storms.

Under the canopy: disentangling the role of stemflow in shaping spatial patterns of soil microbial community structure underneath trees.

Stemflow is a spatially concentrated input of rainwater at the base of trees, resulting from precipitation draining down tree branches to the stem. Depending on tree shape, stemflow can represent a significant fraction of the total rainfall that contacts the tree's canopy area, and can become chemically enriched along its drainage path. As a result, stemflow has been hypothesized to influence microbial communities in the receiving soil proximal to the stem. However, previous studies have (i) yielded conflicting results on the significance of stemflow as a driver in bacterial community composition, and (ii) not directly compared communities in soils with and without stemflow receipt. In this study, a stemflow diversion system was employed on Quercus virginiana trees in Skidaway Island (Georgia, USA) to directly compare soil bacterial communities receiving no stemflow to those beneath trees with no diversion system in place. In both treatments, sample distance from the stem significantly influenced bacterial community structure. However, the absence of stemflow resulted in increased bacterial community diversity across all samples. Stemflow diversion also significantly altered longitudinal patterns in the abundance of multiple taxonomic groups. These results support the hypothesis that Q. virginiana stemflow has a significant impact on bacterial soil inhabitants and is a key factor in taxon selection in stem-proximal communities.

Calcium and aluminum cycling in a temperate broadleaved deciduous forest of the eastern USA: relative impacts of tree species, canopy state, and flux type.

Ca/Al molar ratios are commonly used to assess the extent of aluminum stress in forests. This is among the first studies to quantify Ca/Al molar ratios for stemflow. Ca/Al molar ratios in bulk precipitation, throughfall, stemflow, litter leachate, near-trunk soil solution, and soil water were quantified for a deciduous forest in northeastern MD, USA. Data were collected over a 3-year period. The Ca/Al molar ratios in this study were above the threshold for aluminum stress (<1). Fagus grandifolia Ehrh. (American beech) had a median annual stemflow Ca/Al molar ratio of 15.7, with the leafed and leafless values of 12.4 and 19.2, respectively. The corresponding Ca/Al molar ratios for Liriodendron tulipifera L. (yellow poplar) were 11.9 at the annual time scale and 11.9 and 13.6 for leafed and leafless periods, respectively. Bayesian statistical analysis showed no significant effect of canopy state (leafed, leafless) on Ca/Al molar ratios. DOC was consistently an important predictor of calcium, aluminum, and Ca/Al ratios. pH was occasionally an important predictor of calcium and aluminum concentrations, but was not a good predictor of Ca/Al ratio in any of the best-fit models (of >500 examined). This study supplies new data on Ca/Al molar ratios for stemflow from two common deciduous tree species. Future work should examine Ca/Al molar ratios in stemflow of other species and examine both inorganic and organic aluminum species to better gauge the potential for, and understand the dynamics of, aluminum toxicity in the proximal area around tree boles.

Meteorological influences on stemflow generation across diameter size classes of two morphologically distinct deciduous species.

Many tree species have been shown to funnel substantial rainfall to their stem base as stemflow flux, given a favorable stand structure and storm conditions. As stemflow is a spatially concentrated flux, prior studies have shown its impact on ecohydrological and biogeochemical processes can be significant. Less work has been performed examining stemflow variability from meteorological conditions compared to canopy structural traits. As such, this study performs multiple regressions: (1) to examine stemflow variability due to event-based rainfall amount, intensity, mean wind speeds, and vapor pressure deficit; (2) across three diameter size classes (10-20, 21-40, and >41 cm DBH); and (3) for two common tree species in the northeastern USA of contrasting canopy morphology--Liriodendron tulipifera L. (yellow poplar) versus Fagus grandifolia Ehrh. (American beech). On the whole, multiple regression results yielded significant positive correlations with stemflow for rainfall amount, intensity, and mean wind speed and a significant negative correlation for vapor pressure deficit (VPD). Tree size altered stemflow-meteorological condition relationships, where larger trees strengthened indirect stemflow-VPD and direct stemflow-rainfall and stemflow-intensity associations. Canopies of rougher bark and lower branch angle (represented by L. tulipifera) enhanced correlations for nearly all meteorological conditions via greater stemflow residence time (and longer exposure to meteorological conditions). Multiple regressions performed on leafless canopy stemflow resulted in an inverse relationship with wind speeds, likely decoupling stemflow sheltered solely on bark surfaces from VPD influences. Leaf presence generally increased direct stemflow associations with rainfall intensity, yet diminished stemflow-rainfall relationships. F. grandifolia canopies (exemplifying structures of smoother bark and greater branch angle) strengthened differences in stemflow associations with rainfall/mean wind speed between leaf states. These findings are placed in a conceptual interception loss path analysis, which shows the potential to alter common interception loss estimates in high stemflow stands.

Drought decreases water storage capacity of two arboreal epiphytes with differing ecohydrological traits.

Arboreal epiphytes, plants that grow on trees, can significantly increase rainwater storage and evaporation (i.e., "interception") within canopies. Drought conditions may affect this hydrological role, as epiphytes' physiological responses change leaf properties that affect water retention. Drought-induced changes in epiphyte water storage capacity could substantially alter canopy hydrology, but have not been studied. We tested the effects of drought on the water storage capacity (Smax) of leaves and leaf properties of two epiphytes with distinct ecohydrological traits: resurrection fern (Pleopeltis polypodioides), and Spanish moss (Tillandsia usneoides). Both species are common in maritime forests of the Southeastern USA, where climate change is expected to decrease precipitation in spring and summer. To simulate drought, we dried leaves to 75 %, 50 %, and ~25 % of fresh weight, and quantified their Smax in fog chambers. We measured relevant leaf properties: hydrophobicity, minimum leaf conductance (gmin; a measure of water loss under drought), and Normalized Difference Vegetative Index (NDVI). We found that drought significantly reduced Smax and increased leaf hydrophobicity for both species, indicating that lower Smax may be due to shedding of droplets. While the overall reduction in Smax did not differ between the two species, they exhibited distinct drought responses. Dehydrated T. usneoides leaves had lower gmin, demonstrating the ability to limit water loss under drought. P. polypodioides increased gmin when dehydrated, consistent with its extraordinary ability to withstand water loss. NDVI decreased with dehydration in T. usneoides but not P. polypodioides. Our results suggest that increased drought may have a dramatic effect on canopy water cycling by reducing the Smax of epiphytes. Reduced rainfall interception and storage in forest canopies could have widespread effects on hydrological cycling, thus understanding the potential feedbacks of plant drought response on hydrology is crucial. This study highlights the importance of connecting foliar-scale plant response with broader hydrological processes.

Rain harvesting by plants.

Biddick and Van Stan II introduce how certain plant species harvest rain.

Mini-Review: Stemflow as a Resource Limitation to Near-Stem Soils.

Stemflow, a precipitation and solute supply to soils near tree stems, can play a wide array of roles in ecosystem functioning. However, stemflow's ecohydrological functions have been primarily studied in forests with voluminous stemflow because resource subsidy is currently considered stemflow's only impact on near-stem soils. This common assumption ignores controls that stemflow generation may exert via resource limitation (when stemflow < open rainfall and near-stem throughfall is negligible). We reviewed selected literature across numerous forests to evaluate the predominance of stemflow as a potential resource limitation to near-stem soils and characterized the concentrated, but meager, solute flux from low stemflow generators. Global observations of stemflow were highly skewed (skewness = 4.6) and leptokurtic (kurtosis = 28.8), where 69% of observations were ≤2% of rainfall. Stemflow ≤ 2% of rainfall is 10-100 times more chemically enriched than open rainfall, yet low volumes result in negligible solute fluxes (under 1 g m-2 y-1). Reduced stemflow may be the global and regional norm, creating persistently dry near-stem soils that receive infrequent, salty, and paltry precipitation flux if throughfall is also low. Ignoring stemflow because it results in scarcity likely limits our understanding of ecosystem functioning as resource limitations alter the fate of soil nutrients, energy flows, and spatial patterning of biogeochemical processes.

A review and evaluation of forest canopy epiphyte roles in the partitioning and chemical alteration of precipitation.

Interactions between precipitation and forest canopy elements (bark, leaves, and epiphytes) control the quantity, spatiotemporal patterning, and the chemical concentration, character and constituency of precipitation to soils. Canopy epiphytes exert a range of hydrological and biogeochemical effects due to their diversity of morphological traits and nutrient acquisition mechanisms. We reviewed and evaluated the state of knowledge regarding epiphyte interactions with precipitation partitioning (into interception loss, throughfall, and stemflow) and the chemical alteration of net precipitation fluxes (throughfall and stemflow). As epiphyte species are quite diverse, this review categorized findings by common paraphyletic groups: lichens, bryophytes, and vascular epiphytes. Of these groups, vascular epiphytes have received the least attention and lichens the most. In general, epiphytes decrease throughfall and stemflow and increase interception loss. Epiphytes alter the spatiotemporal pattern of throughfall and increase overall latent heat fluxes from the canopy. Epiphytes alter biogeochemical processes by impacting the transfer of solutes through the canopy; however, the change in solute concentration varies with epiphyte type and chemical species. We discuss several important knowledge gaps across all epiphyte groups. We also explore innovative methods that currently exist to confront these knowledge gaps and past techniques applied to gain our current understanding. Future research addressing the listed deficiencies will improve our knowledge of epiphyte roles in water and biogeochemical processes coupled within forest canopies-processes crucial to supporting microbe, plant, vertebrate and invertebrate communities within individual epiphytes, epiphyte assemblages, host trees, and even the forest ecosystem as a whole.

The effects of phenoseason and storm characteristics on throughfall solute washoff and leaching dynamics from a temperate deciduous forest canopy.

Seasonal variations in the washoff and leaching dynamics of throughfall ionic fluxes represent a significant process affecting the biogeochemical cycling of forested ecosystems-particularly for temperate deciduous forests with distinct phenological seasons (or "phenoseasons"). Most studies on temperate deciduous forests aggregate seasonal throughfall fluxes to the leafed (growing) and leafless (dormant) periods, yet the phenological conditions controlling seasonality demand finer-scale demarcations that include the transitional phenoseasons (leaf senescence and emergence). To fill these gaps our study examines the washoff and leaching dynamics of Na(+), Mg(2+), K(+), Ca(2+), Cl(-), SO(4)(2-), and NO(3)(-) throughfall derived from bulk and sequentially sampled rain events across leafed, leafless and both transitional phenoseasons over a 3-year period (2008-2010). As throughfall washoff and leached solute fluxes are also closely-coupled to rainfall conditions, we further examine the effects of storm characteristics on phenoseasonal washoff-dominated (Na(+) and Cl(-)) and leaching-dominated (K(+), Ca(2+), Mg(2+)) fluxes through intrastorm event comparison plots and factorial MANOVA. Highly significant differences in leached and washoff solute fluxes were found across meteorological conditions (p<0.001) nested within phenoseasonal divisions (p<0.00001). Phenoseasonal washoff Na(+) and Cl(-) fluxes seemed to be more closely related to leaf area; whereas, leaching flux and canopy exchange of all solutes to correspond more with major phenological changes (when the canopies tend to be most metabolically active). The greatest differences in leached Mg(2+), K(+), Ca(2+), and SO(4)(2-) fluxes were not between the full leafed and leafless phenoseasons (33-80% difference), but between the transitional periods (80 to 200 fold greater during leaf senescence than leaf emergence). Intrastorm average canopy NO(3)(-) leaching, however, ranged from low losses (1 µmol(c)m(-2)h(-1)) to canopy uptake (-2 µmol(c)m(-2)h(-1)) during both transitional phenoseasons. K(+), Ca(2+), Mg(2+) were all markedly more exchangeable during senescence, with Ca(2+) and Mg(2+) being more tightly held by the canopy. Leaching rates and fluxes for all measured solutes were negligible to negative during emergence, except for K(+) and SO(4)(2-). Our results indicate that much of the variance in timing and magnitude of throughfall solute fluxes to forest soils within temperate deciduous ecosystems may be ascribed to phenologically-delineated seasons and storm conditions.