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  1. Abstract

    Water in rivers is delivered via the critical zone (CZ)—the living skin of the Earth, extending from the top of the vegetation canopy through the soil and down to fresh bedrock and the bottom of significantly active groundwater. Consequently, the success of stream‐rearing salmonids depends on the structure and resulting water storage and release processes of this zone. Physical processes below the land surface (the subsurface component of the CZ) ultimately determine how landscapes “filter” climate to manifest ecologically significant streamflow and temperature regimes. Subsurface water storage capacity of the CZ has emerged as a key hydrologic variable that integrates many of these subsurface processes, helping to explain flow regimes and terrestrial plant community composition. Here, we investigate how subsurface storage controls flow, temperature, and energetic regimes that matter for salmonids. We illustrate the explanatory power of broadly applicable, storage‐based frameworks across a lithological gradient that spans the Eel River watershed of California. Study sites are climatically similar but differ in their geologies and consequent subsurface CZ structure that dictates water storage dynamics, leading to dramatically different hydrographs, temperature, and riparian regimes—with consequences for every aspect of salmonid life history. Lithological controls on the development of key subsurface CZ properties like storage capacity suggest a heretofore unexplored link between salmonids and geology, adding to a rich literature that highlights various fluvial and geomorphic influences on salmonid diversity and distribution. Rapidly advancing methods for estimating and observing subsurface water storage dynamics at large scales present new opportunities for more clearly identifying landscape features that constrain the distributions and abundances of organisms, including salmonids, at watershed scales.

     
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  2. Abstract

    Bedrock vadose zone water storage (i.e., rock moisture) dynamics are rarely observed but potentially key to understanding drought responses. Exploiting a borehole network at a Mediterranean blue oak savanna site—Rancho Venada—we document how water storage capacity in deeply weathered bedrock profiles regulates woody plant water availability and groundwater recharge. The site is in the Northern California Coast Range within steeply dipping turbidites. In a wet year (water year 2019; 647 mm of precipitation), rock moisture was quickly replenished to a characteristic storage capacity, recharging groundwater that emerged at springs to generate streamflow. In the subsequent rainless summer growing season, rock moisture was depleted by about 93 mm. In two drought years that followed (212 and 121 mm of precipitation) the total amount of rock moisture gained each winter was about 54 and 20 mm, respectively, and declines were documented exceeding these amounts, resulting in progressively lower rock moisture content. Oaks, which are rooted into bedrock, demonstrated signs of water stress in drought, including reduced transpiration rates and extremely low water potentials. In the 2020–2021 drought, precipitation did not exceed storage capacity, resulting in variable belowground water storage, increased plant water stress, and no recharge or runoff. Rock moisture deficits (rather than soil moisture deficits) explain these responses.

     
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  3. Abstract

    Cross‐slope climate differences in the midlatitudes are ecologically important, and impact vegetation‐mediated water balance between the earth surface and the atmosphere. We made high‐resolution in situ observations of air temperature, relative humidity, soil moisture, insolation, and sap velocity observations on 14 Pacific madrone trees (Arbutus menziesii) spanning adjacent north and south slopes at the University of California's Angelo Coast Range Reserve. To understand the cross‐slope response of sap velocity, a proxy for transpiration, to microclimate, we modeled the sap velocity on each slope using a transpiration model driven by ambient environment and parameterized with a Markov Chain Monte Carlo parameter estimation process. The results show that trees on opposing slopes do not follow a shared pattern of physiological response to transpiration drivers. This means that the observed sap velocity differences are not due entirely to observed microclimate differences, but also due to population‐level physiological differences, which indicates acclimation to inhabited microclimate. While our present data set and analytical tools do not identify mechanisms of acclimation, we speculate that differing proportions of sun‐adapted and shade‐adapted leaves, differences in stomatal regulation, and cross‐slope root zone moisture differences could explain some of the observed and modeled differences.

     
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  4. Abstract

    The relative importance of separation by distance and by environment to population genetic diversity can be conveniently tested in river networks, where these two drivers are often independently distributed over space. To evaluate the importance of dispersal and environmental conditions in shaping microbial population structures, we performed genome‐resolved metagenomic analyses of benthicMicrocoleus‐dominated cyanobacterial mats collected in the Eel and Russian River networks (California, USA). The 64 Microcoleusgenomes were clustered into three species that shared >96.5% average nucleotide identity (ANI). Most mats were dominated by one strain, but minor alleles within mats were often shared, even over large spatial distances (>300 km). Within the most commonMicrocoleusspecies, the ANI between the dominant strains within mats decreased with increasing spatial separation. However, over shorter spatial distances (tens of kilometres), mats from different subwatersheds had lower ANI than mats from the same subwatershed, suggesting that at shorter spatial distances environmental differences between subwatersheds in factors like canopy cover, conductivity, and mean annual temperature decreases ANI. Since mats in smaller creeks had similar levels of nucleotide diversity (π) as mats in larger downstream subwatersheds, within‐mat genetic diversity does not appear to depend on the downstream accumulation of upstream‐derived strains. The four‐gamete test and sequence length bias suggest recombination occurs between almost all strains within each species, even between populations separated by large distances or living in different habitats. Overall, our results show that, despite some isolation by distance and environmental conditions, sufficient gene‐flow occurs among cyanobacterial strains to prevent either driver from producing distinctive population structures across the watershed.

     
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  5. Abstract

    Bedrock weathering regulates nutrient mobilization, water storage, and soil production. Relative to the mobile soil layer, little is known about the relationship between topography and bedrock weathering. Here, we identify a common pattern of weathering and water storage across a sequence of three ridges and valleys in the sedimentary Great Valley Sequence in Northern California that share a tectonic and climate history. Deep drilling, downhole logging, and characterization of chemistry and porosity reveal two weathering fronts. The shallower front is ∼7 m deep at the ridge of all three hillslopes, and marks the extent of pervasive fracturing and oxidation of pyrite and organic carbon. A deeper weathering front marks the extent of open fractures and discoloration. This front is 11 m deep under two ridges of similar ridge‐valley spacing, but 17.5 m deep under a ridge with nearly twice the ridge‐valley spacing. Hence, at ridge tops, the fraction of the hillslope relief that is weathered scales with hillslope length. In all three hillslopes, below this second weathering front, closed fractures and unweathered bedrock extend about one‐half the hilltop elevation above the adjacent channels. Neutron probe surveys reveal that seasonally dynamic moisture is stored to approximately the same depth as the shallow weathering front. Under the channels that bound our study hillslopes, the two weathering fronts coincide and occur within centimeters of the ground surface. Our findings provide evidence for feedbacks between erosion and weathering in mountainous landscapes that result in systematic subsurface structuring and water routing.

     
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  6. Abstract

    Duration and temporal stability of resource subsidy largely affect the response of recipient communities. Factors that influence the temporal dynamics of resource subsidy from aquatic‐to‐terrestrial habitats by emerging aquatic insects were examined in this study. By measuring the flux of aquatic insect emergence from six habitats in a river over summer, we found that the timing of emergence varied by habitats for each dominant taxa, and that different species emerged at different times of the summer sequentially. We found that spatial variation in the emergence timing caused by the spatial heterogeneity of the water temperature, and so on in the source habitat can temporally stabilize the subsidy of each species from the whole river. Similarly, we found that the variation in emergence timing between species contributed to the temporal stability of subsidies from each habitat. The contribution of spatial heterogeneity to the temporal stability varied by the focal species and the contribution of species diversity varied by habitats. This study demonstrates how the ecological function of spatial heterogeneity and species diversity crosses the boundary of ecosystems by temporally stabilizing resource subsidies.

     
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  7. Abstract

    Soils are widely considered the primary terrestrial organic matter pool mediating carbon transactions with the atmosphere and groundwater. Because soils are both a host and a product of rhizosphere activity, they are thought to mark the location where photosynthetic fixation of carbon dioxide (CO2) is balanced by the oxidation of organic matter. However, in many terrestrial environments, the rhizosphere extends below soils and into fractured bedrock, and it is unknown if the resulting biological and hydrologic dynamics in bedrock have a significant impact on carbon cycling. Here we show substantial production of CO2in weathered bedrock at 4–8 m below the thin soils (<0.5 m thick) of a Northern California forest using innovative monitoring technology for sampling gases and water in fractured rock. The deep CO2production supports a persistent upward flux of CO2(g)year‐round from bedrock to soil, constituting between 2% and 29% of the average daily CO2efflux from the land surface. When water is rapidly transported across the fractured bedrock vadose zone, nearly 50% of the CO2produced in the bedrock dissolves into water, promoting water‐rock interaction and export of dissolved inorganic carbon (DIC) from the unsaturated zone to groundwater, constituting as much as 80% of the DIC exiting the hillslope. Such CO2production in weathered bedrock is subject to unique moisture, temperature, biological, and mineralogical conditions which are currently missing from terrestrial carbon models.

     
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  8. Abstract

    The spatiotemporal dynamics of plant water sources are hidden and poorly understood. We document water source use ofQuercus garryanagrowing in Northern California on a profile of approximately 50 cm of soil underlain by 2–4 m of weathered bedrock (sheared shale mélange) that completely saturates in winter, when the oaks lack leaves, and progressively dries over the summer. We determined oak water sources by combining observations of water stable isotope composition, vadose zone moisture and groundwater dynamics, and metrics of tree water status (potential) and use (sapflow). During the spring, oak xylem water is isotopically similar to the seasonal groundwater and shallow, evaporatively enriched soil moisture pools. However, as soils dry and the water table recedes to the permanently saturated, anoxic, low‐conductivity fresh bedrock boundary,Q. garryanashifts to using a water source with a depleted isotopic composition that matches residual moisture in the deep soil and underlying weathered bedrock vadose zone. Sapflow rates remain high as late‐summer predawn water potentials drop below2.5 MPa. Neutron probe surveys reveal late‐summer rock moisture declines under the oaks in contrast to constant rock moisture levels under grass‐dominated areas. We therefore conclude that the oaks temporarily use seasonal groundwater when it occupies the weathered profile but otherwise use deep unsaturated zone moisture after seasonal groundwater recedes. The ample moisture, connected porosity, and oxygenated conditions of the weathered bedrock vadose zone make it a key tree water resource during the long summer dry season of the local Mediterranean climate.

     
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  9. Summary

    The emergence of critical zone (CZ) science has provided an integrative platform for investigating plant ecophysiology in the context of landscape evolution, weathering and hydrology. The CZ lies between the top of the vegetation canopy and fresh, chemically unaltered bedrock and plays a pivotal role in sustaining life. We consider what the CZ perspective has recently brought to the study of plant ecophysiology. We specifically highlight novel research demonstrating the importance of the deeper subsurface for plant water and nutrient relations. We also point to knowledge gaps and research opportunities, emphasising, in particular, greater focus on the roles of deep, nonsoil resources and how those resources influence and coevolve with plants as a frontier of plant ecophysiological research.

     
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  10. Abstract

    Despite a multitude of small catchment studies, we lack a deep understanding of how variations in critical zone architecture lead to variations in hydrologic states and fluxes. This study characterizes hydrologic dynamics of 15 catchments of the U.S. Critical Zone Observatory (CZO) network where we hypothesized that our understanding of subsurface structure would illuminate patterns of hydrologic partitioning. The CZOs collect data sets that characterize the physical, chemical, and biological architecture of the subsurface, while also monitoring hydrologic fluxes such as streamflow, precipitation, and evapotranspiration. For the first time, we collate time series of hydrologic variables across the CZO network and begin the process of examining hydrologic signatures across sites. We find that catchments with low baseflow indices and high runoff sensitivity to storage receive most of their precipitation as rain and contain clay‐rich regolith profiles, prominent argillic horizons, and/or anthropogenic modifications. In contrast, sites with high baseflow indices and low runoff sensitivity to storage receive the majority of precipitation as snow and have more permeable regolith profiles. The seasonal variability of water balance components is a key control on the dynamic range of hydraulically connected water in the critical zone. These findings lead us to posit that water balance partitioning and streamflow hydraulics are linked through the coevolution of critical zone architecture but that much work remains to parse these controls out quantitatively.

     
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