HYDROLOGICAL PROCESSES Hydro/. Process. (2011) Published online in Wiley Online Library (wileyonlinelibrary.com). 001: 10.1002jhyp.8119

The role of the hyporheic zone across stream networks t

Steven M. Wondzell* Abstract

USDA Forest Service, Pacific Many hyporheic papers state that the hyporheic zone is a critical component of Northwest Research Station, Olympia stream ecosystems, and many of these papers focus on the biogeochemical effects Forestry Sciences Laboratory, of the hyporheic zone on stream solute loads. However, efforts to show such Olympia, WA 98512, USA relationships have proven elusive, prompting several questions: Are the effects of the hyporheic zone on stream ecosystems so highly variable in place and time *Correspondence to: (or among streams) that a consistent relationship should not be expected? Or, is the Steven M. Wondzell, USDA Forest hyporheic zone less important in stream ecosystems than is commonly expected? Service, Pacific Northwest Research These questions were examined using data from existing modelling Station, Olympia Forestry Sciences Laboratory, Olympia, WA studies of hyporheic exchange flow at five sites in a fifth-order, mountainous stream 98512, USA. network. The size of exchange flows, relative to stream discharge (QHEF : Q), was E-mail: [email protected] large only in very small streams at low discharge (area ",,100 ha; Q <10I/s). At higher flows (flow exceedance probability >0·7) and in all larger streams, was small. These data show that biogeochemical processes in the +This article is a US Government work QHEF : Q and is in the public domain in the USA. hyporheic zone of small streams can substantially influence the stream's solute load, but these processes become hydrologically constrained at high discharge or in larger streams and . The hyporheic zone may influence stream ecosystems in many ways, however, not just through biogeochemical processes that alter stream solute loads. For example, the hyporheic zone represents a unique habitat for some , with patterns and amounts of and downwelling water determining the underlying physiochemical environment of the hyporheic zone. Similarly, hyporheic exchange creates distinct patches of downwelling and upwelling. Upwelling environments are of special interest, because upwelling water has the potential to be thermally or chemically distinct from stream water. Consequently, micro-environmental patches created by hyporheic exchange flows are likely to be important to biological and ecosystem processes, even if their impact on stream solute loads is small. Published in 2011 by John Wiley & Sons, Ltd.

Key Words hyporheic exchange flows; stream discharge; stream networks; flow exceedance probability; watershed area; hyporheic potential

Introduction There is growing recognition that the part of streams extending below, and adjacent to, the streambed can be an important component of aquatic ecosys­ tems. This area, commonly known as the hyporheic zone, can provide unique habitats for aquatic organisms (Stanford and Ward, 1988; Baxter and Hauer, 2000), and exchange of stream water through the hyporheic zone exposes transported solutes to unique biogeochemical environments with subsequent impacts on whole stream metabolism (Grimm and Fisher, 1984) and nutri­ ent cycling (Triska et al., 1989; Mulholland et aI., 1997; Mulholland and DeAngelis, 2000). Despite its broadly recognized importance, quantifying the role of the hyporheic zone in streams has proven difficult. For example, attempts to examine this question in studies comparing a wide range of stream types often do not show strong correlations between measures of hyporheic exchange and nutrient cycling at the stream-reach scale (Hall et aI., 2002; Webster et al., 2003). These results pose two questions: (i) Are the effects of the hyporheic zone on stream ecosystems so highly variable in place and time Received 15 August 2010 (or among streams) that a consistent relationship should not be expected? Or Accepted 30 March 2011 (ii), is the hyporheic zone less important in stream ecosystems than is com­ monly expected? To answer these questions, I take a physical/hydrological

Published in 2011 by John Wiley & Sons, ltd. S. M. WONDZEU.

Table L Data summlliy for points shown in Figure 1. Abbreviations shown in the first column were used in Figure I to denote specific stTeam reaches from the 64 km2 Lookout Creek watershed in central Oo·egon. USA. Slope denotes [he longitudinal gradiem 8f the valley floor; QHEF denotes the amount of hyporheic exchange flow predicted from groundwater flow models. normalized to a lOO-m ,·each length; Ex (p) denotes the annual flow exceedance probability for a given stream under the flow conditions used to simulate hyporheic exchange flows in the groundwater flow models. Data sources are as follows: WS1 and \VS3 (Kasahara and \Vondzeli. 2003) with additional comparison for low- and high-baseftow corrditions (Wondzel1. 2006); McRae Creek (Wondzell and Swanson. 1996): upper and lower Lookout Creek (Kasahara and Wondzell. 2003)

Site \Vatershed Area Mean Slope K HYPpor QHH Q QHf,IQ Ex annual (Slope x -1 1 .l. (p) (Inin�l) (m s- ') K; S per S- ) (n1- ) -l discharge m S ) 100 m) (lIs) l-L WS1�low baseflow 0·96 40 0·14 7x 10-5 1 X 10-5 2·28 ]·22 0·01869 0·90 l-H WS i �high baseftow 0·96 40 0·14 7 x 10-5 1 X 10-5 2·28 4·67 0·00488 0·69 3-L WS3�low baseflow j·O l 39 o 13 7x 10-5 9x 10-0 4·26 3·23 0·01316 0-92 3-H WS3�high base flow 101 39 o 13 7x 10-) 9 X 10-6 4·26 1146 0·00372 0·61 McR-L McRae�low baseflow 14·00 530 0·04 5 x 10-' LX 10-4 ]·27 117 0·0001! 0·71 McR-H McRae�high baseflow 14·00 530 0·04 5x ]0-' 2x 10-4 175 590 0·00003 0·30 McR-S McRae�storm flow 14·00 530 0·04 5x 10-' 2x 10-'; 149 2400 0·00001 0·03 UpLO upper Lookout�basefiow 50 00 2630 002 Ix 10-' 3 X 10-5 50·0 308 0·00069 0·86 LowLO lower Lookout�baseflow 60·62 3190 0·01 7x 1,,-4v 7 X 10-6 l14 873 0·00002 0·67 perspective and attempt to quantify the importance of each study site were estimated from simulations using the the hyporheic zone in a mountainous stream network. numerical groundwater flowmodel, MODFLOW. Models From these results, I build a general conceptual model were parameterized to measured stream boundary condi­ to describe the expected role of the hyporheic zone in tions and hydraulic conductivi�ies measured from stream networks. tests from relatively dense well networks covering the full model domains. Models were calibrated to simulate An example from the Lookout Creek watershed the head distributions measured across the well networks. In-depth study at one study site suggested groundwater Lookout Creek drains a 64 km2 mountainous water­ flow models provide reasonable quantitative estimates of shed located in the western Cascade Mountains of Ore­ hyporheic exchange fluxes (Wondzell et ai .. 2009a). gon, USA (44°20'N. 122'20!W). Hyporheic studies and Results from the Lookout Creek watershed showed groundwater flow simulations were conducted at five that QHEF: Q ratios decreased as stream size increased. locations within the watershed. Two sites were located Because the QHEF : Q ratio is sensitive to change in dis­ on the fifth-order mainstem of Lookout Creek. The lower charge. comparisons among streams of different sizes Lookout site was located in a narrow. bedrock-confined within a stream network may be confounded changes reach whereas the upper Lookout site was located in in Q if the observations are made at different times and a wide. or unconstrained alluvial reach. One site was under different flow conditions. One way to control for located on McRae Creek. a fourth-order tributary to changes in discharge is to make among-site comparisons Lookout C;eek. Two sites were located on first-order at discharges with similar flow exceedance probabilities. headwater streams, both of which drained directly to Within a smali watershed, or even within a hydro-climatic lower Lookout Creek (Table I). These study sites are region, the flow exceedance probability provides seme all located in forested watersheds where road build­ insurance that comparisons are made under similar hydro­ ing and forest harvest have occurred in the past. Large logic conditions. In this case, calibration data for model wood was also removed from portions of the stream simulations at a number of sites in the Lookout Creek network, including much of the fifth-order stream chan­ basin were collected around 0·70 flow exceedance prob­ nel. Much of the road network remains in and abilities, providing reasonably comparable data across a in use. More detailed study site descriptions and the wide range of stream sizes (Table 0. These data showed specifics of the model simulations are in Wondzell relatively high QHEF: Q ratios in the headwater WS1 and (1994). Wondzell and Swanson (1996), and Kasat'1ara and WS3 streams and much smailer ratios in the larger McRae Wondzell (2003). Creek and Lookout Creek sites. The relationship between hyporheic exchange flow and Results from the Lookout Creek watershed also stream discharge at the five study sites in the Lookout showed that QHEF: Q ratios decreased with seasonal Creek watershed was examined for systematic patterns increases in discharge. For example, at McRae Creek, related either to temporal changes in discharge at a single only small changes in hyporheic exchange flows (QHEF) site or changes across a stream network under relatively were observed over a 20-fold change in stream dis­ uniform flow conditions. Hyporheic exchange flows at charge from summer low baseflow through winter high

Published in 2011 by John Wiley & Sons, Ltd. Hydro/. Process. (2011) SCIENTIFIC BRIEFING baseflow-to-peak flows during a smaii storm (Table I). reach as described by Darcy's Law (Wondzell and Consequently. the QHEF : Q ratio decreased approximately Gooseff, in press). Head gradients created by changes 15-fold as discharge increased. Hyporheic exchange was in the longiwdina! gradients along stream channels are only simulated for low baseftow conditions at WS 1 and a major factor driving hyporheic exchange in many WS3 (Kasahara and WondzelL 2003). However, direct streams. Where average longitudinal gradients are steep, observations from the wei! networks at these sites showed steep head gradients can develop over short distances that hyporheic exchange fluxes should have remained across steps, riffles and channel meanders: where gra­ relatively constant over the observed range of base­ dients are low. long distances are required to create sub­ flow discharges because neither the shape of the water stantial head differences so that head gradients also tend table nor stream water elevations changed substantially. to be low. Of course, in low-gradient streams the head Consequently, the QHEF : Q ratios would have decreased loss along the longitudinal profile can be compressed into markedly with relatively modest increases in discharge distinct steps but steps or riffles must then be widely (Table I). spaced. Consequently, valleys with steeper longitudinal While the general trend of decreasing QHEF:Q with gradients can support steeper head gradients. resulting in increasing discharge is readily apparent from the Lookout more hyporheic exchange. Longitudinal valley gradients Creek data. it is also clear that there is substantial reach to tend to change systematically with stream size in many reach variation not explained by simple differencesin dis­ mountainous stream networks such that small headwater charge (Table I). For example, the two sites on the main­ channels tend to have steeper longitudinal gradients than stem of Lookout Creek have roughly similar discharges. do larger streams. These trends are clearly evident in the but QHEF is more than lOO-fold greater at upper Look­ data collected from the Lookout Creek stream network out Creek than at lower Lookout Creek. Lower Look­ (Table I). out Creek flows through a bedrock-constrained reach in Darcy's Law also shows that hyporheic exchange which hyporheic exchange is limited to a narrow gravel should also be linearly rdated to saturated hydraulic bar on one side of the channel with the other side of conductivity (K), with more exchange flow through the channel pressed against exposed bedrock. In contrast, coarse textured sediment with high K and less flow the upper Lookout Creek study site is located in one through fine textured sediment with low K. There is a of the widest reaches found along the length of main­ tendency for fining of alluvial sediment with decreased stem Lookout Creek and extensive hyporheic exchange longitudinal gradient, and gradient tends to decrease occurs throughout the reach. In this case. differences with watershed area, as described above. Consequently, in channel morphology led to a 40-fold change in the high-gradient mountain streams would tend to have QHEF: Q ratio-a change larger than that observed at coarse streambed sediment and valley floor alluvium McRae Creek from low baseflow to storm flow (Table I). whereas lower gradient lowland stream networks and The QHEF : Q ratios at low flowwere also somewhat dif­ mainstems of major rivers would tend to be characterized ferent between the WS 1 and the WS3 study sites because by finer sediment. Of course, these patterns are also of differences in floor widths and the abundance under substantial local control. There is not a consistent of log jams (Wondzell, 2006). pattern in K with stream size in the Lookout Creek stream network (Table I). All the study sites tend to A Generalized Conceptual Model be relatively high gradient with correspondingly high of the Hyporheic Zone average Ks, typically in the range observed for gravels to sands (Domenico and Schwartz, 1990, Table m.2, The analyses presented above suggest that the relative pg. 65). size of the hyporheic zone (QHEF: Q ratio) is inversely Multiplying longitudinal gradient of the valley proportional to stream discharge. This relationship also by the saturated hydraulic conductivity measured in any appears sensitive to the underlying potential for a given stream reach provides an initial estimate of the potential stream reach to support hyporheic exchange. Thus, broad­ to develop hyporheic exchange (HYPPOT). This product scaie, among site comparisons need to account for both is roughly analogous to the Darcy Velocity, but uses factors. The discharge regime is easily characterized by reach-scale longitudinal gradient of the valley floor rather simple metrics such as watershed area, annual average than actual estimates of (.6.h/.6.1). Expressed L1is way, discharge, and flow exceedance probability. Characteris­ HYPPOT could exceed 10-3 m S-1 in mountain streams ing the potential of a stream reach to support hyporheic where longitudinal gradients of alluvial reaches can equal exchange is more difficult. 0· 15 mm-i and where K in coarse valley-floor alluvium might be similar to that of fine gravels. At the other The potential for hyporheic exchange extreme, HYPPOT could be as low as 10-15 m S-I, in The potential for hyporheic exchange will be related to low-gradient streams in or tidal mud fiats both head gradients (.6.h/.6.1) and the saturated hydraulic that can have gradients less than 10-4 m'm-1 and K conductivity (K) of alluvial sediment within a stream typical of clayey sediment. Applying the HYPPOT metric

Published in 2011 by John Wiley & Sons, Ltd. Hydro/. Process. (2011) S. M. WONDZELL

Ie - ". ""fu "x

FlOw exceedOnce prObabil�ies c:=:J 0.90 c:=:J 0.70 c:=:JO,5(} c:::J.0.30

Figure L Conceptual depiction of the relative influence of the hyporheic zone on bulk stream water (the ratio of QHEF/Q) relative to watershed area and underlying potential of the stream network to support hyporheic exchange flow. Data points are from the Lookout Creek watershed and point labels follow Table L The 3-D response surfaces are hypothesized relationships showing how the of QHEF/Q ratio might be expected to change with change in watershed area and the hydrogeomorphic conditions that create the potential for hyporheic exchange (HYPPOT). Each surface represents flow conditions characterized by the annual flow exceedance probability. from periods of very low flow (90% exceedance) to periods of moderately high flow (30% exceedance) to the step-pool and pool-riffle stream reaches of the exchange relative to stream discharge, nor do they pro­ mountainous Lookout Creek network suggests that the vide quantitative comparisons between the head gradients potential for hyporheic exchange flow is high, which generated by pumping exchange and longitudinal valley agrees well with the high QHEF: Q ratios estimated for gradients in streams characterized by dune-ripple channel these stream reaches. Similarly, the HYPPOT metric is morphology. Thus, it is currently l not possible to evalu­ likely to work well in meandering lower-gradient streams ate how well HYPPOT would characterize the potential because both valley slope and hydraulic conductivity for pumping exchange. However, that the streambed sed­ are key variables in the equations shown by Cardenas iment is fine textured suggests that the magnitude of (2009) to predict hyporheic exchange in meandering hyporheic exchange is likely to be limited by relatively rivers. low K, a relationship which is clearly captured by the The HYPPOT metric does not account for all the fac­ HYPpoT metric. Thus, despite some limitation, the simple tors known to drive hyporheic exchange. Reach-scale HYPpoT metric should provide an initial rough estimate changes in channel morphology can lead to large changes of the potential for hyporheic exchange within a given stream reach and can therefore help facilitate among-site in QHEF, even though K and longitudinal gradients remain comparisons. unchanged. For example, HYPpoT does not distinguish between the lower and upper Lookout Creek sites where differences in channel constraint result in more than a Comparison among streams lO-fold difference in QHEF between these sites (Table I). The expected behaviour of the hyporheic zone across The HYPPOT metric is not sensitive to transient exchange a wide range of stream types and sizes can be com­ driven by rapid changes in stage, such as in tidally pared using the QHEF: Q ratio, stream size as measured influenced rivers (Bianchin et al., 2010). Perhaps more by watershed area or average annual discharge, flow importantly, HYPPOT is not directly sensitive to the inter­ exceedance probability, and HYPpoT (Figure 1). actions between flow and stream bedforms that control The QHEF : Q ratio is large only in small streams, and pumping exchange (Elliott and Brooks, 1997) which is even there, only at relatively low discharges. For exam­ expected to be the dominant form of hyporheic exchange ple, in the smallest streams (W S I & WS3) at late sum­ in many low gradient streams with relatively mobile and mer low flows (exceedance probability �0·9), turnover fine-textured bed sediment. Although pumping exchange lengths of stream water through the hyporheic zone are has been widely studied, published studies do not pro­ as short as 50-75 m. That is, on average, the entire vide quantitative estimates of the magnitude of hyporheic in-channel flow seeps into the hyporheic zone and is

Published in 2011 by John Wiley & Sons, Ltd. Hydrol. Process. (2011) SCIENTIFIC BRIEFING replaced by upwelling hyporheic water over distances as were 8 km in a second-order tributary with summer base­ I short as 50 m. Even at high baseflow (exceedance prob­ flow discharge of only 9 i S- , and 1300 km in the fifth­ ability �0·6-0· 7), the turnover length of stream water order Silver Creek with summer baseftow discharge of l is only 250 m. Thus, these streams are characterized by 344 I S- . These studies appear to have focused only large QHEF: Q ratios so that physical and biogeochemi­ on the specific features where high rates of hyporheic cal processes like nutrient cycling and water temperature exchange were expected. thus, the QHEF: Q ratio is likely regimes would be strongly influencedby interactions with to be even smaller for these streams as a whole. Lacking the hyporheic zone over large parts of the year. These estimates of watershed areas and the flow exceedance small headwater streams make up more than two-thirds probabilities, these data cannot be added to Figure I. of the total stream network within the Lookout Creek However, these observations are consistent with the shape watershed (Wondzell. 1994) and are likely to regulate of the baseflow response surfaces for streams with low solute transport from the entire watershed for large parts HYPpoT which show the QHEF: Q ratios are small for of the year. small streams and decrease to near zero as watershed Lower in the watershed, estimated turnover lengths area increases. range from 0·62 km for upper Lookout Creek (flow There will, of course, be exceptions to the general exceedances probability �0·86) to 77 km for lower Look­ trends described above, with the most notable cases out Creek during summer baseflow (flow exceedances from large, cobble, and gravel-bedded rivers flowing probability �0·67). The wide. unconstrained reach of through unconstrained alluvial valleys in mountainous upper Lookout Creek has a relatively high QHEF: Q ratio regions. Perhaps one of the best known examples is and could be a hot-spot for nutrient processing in the the Nyack Floodplain on the Middle Fork Flathead mainstem stream. However, most of Lookout Creek is River (Poole et ai., 2006) where hyporheic exchange moderately to highly constrained, more similar to the has long been studied. Quantitative estimates of QHEF lower Lookout Creek site, so that the hyporheic zone is have not been published for this site. Synoptic discharge unlikely to strongly influence the mass flux of nutrients measurements, however, suggest a net loss of 30% of transported through Lookout Creek in most locations. the stream discharge to the subsurface in the upper The QHEF: Q ratio decreases as stream discharge Nyack floodplain (Stanford et ai., 1994) suggesting that increases in wet seasons or during storms because QHEF hyporheic exchange might be substantial in this large should decrease as lateral inflows from adjacent hiils river. Similar observations have been made in other begin to restrict the size of the hyporheic zone and reverse unconstrained alluvial river reaches (Laenen and Bencala, head gradients forcing flow across the floodplain toward 200]; Konrad el at., 2005). Unfortunately, published quantitative estimates of hyporheic exchange flows in the stream as shown by Storey et al. (2003). At the large rivers are rare. same time, discharge would increase markedly. At high flows (flow exceedance probabilities >0·3), the turnover lengths of stream water in WS 1 and WS3 would exceed Mass Transport is Not AU That Matters the length of these tributaries. In large streams, lower The dominance of tracer-based approaches used to study in the network, turnover lengths would be even longer the physical of the hyporheic zone, and so that neither stream water nor accompanying solutes associated tracer-based studies of nutrient spiraling in would be affected by processing in the hyporheic zone. streams, has tended to promote a stream-centric view of In periods of high flow, �30% of the year, the hyporheic the hyporheic zone with a focus on solute transport. The zone will have only a minor effect on whole watershed impression that the relative importance of the hyporheic nutrient processing. zone to stream ecosystem processes is determined by the Not all stream types have an equal potential for proportion of the stream discharge flowing through the hyporheic exchange. High gradient mountain streams hyporheic zone (Findlay, 1995; Jones and Holmes, 1996) flowing over coarse alluvium are likely to have among likely stems from this stream-centric view point. The the highest potential for hyporheic exchange. In con­ hyporheic zone has been recognized for its importance trast, hyporheic exchange is likely to be much smaller to stream ecosystems on many fronts, however, so in low gradient streams. Only a few studies have quanti­ that attempts to assess the relative importance of the fied hyporheic exchange flow in lowland streams. Storey hyporheic zone should be broader than simply questions et al. (2003) focused on single riffle where channel mor­ of mass transport. phology is conducive to hyporheic exchange flow. Even so, the QHEF: Q ratio was low at summer baseflow, giv­ Hyporheic exchange creates unique subsurface ing an estimated turnover length of 21 km when stream habitats I discharge was only 100 ] S- . Kasahara and Hill (2006) The hyporheic zone represents a unique habitat for some focused on riffles constructed for stream restoration so organisms, with patterns and amounts of upwelling and that head gradients and K were artificially increased. downwelling water determining the underlying phys­ Even under these conditions, estimated turnover lengths iochemical environment of the hyporheic zone. These

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N t

. - Active channel

t:::::=J Wetted channel ....a Downwelling '" Upwelling --.. Large wood

Direction of flow Scale (m) • o 10 20

Figure 2. Plan view map of the wetted channel of Bambi Creek. Alaska (from Wondzell et a/.. 2009b). Shaded shows zones of downwelling and upwelling located on streambed and fine-dashed arrows qualitatively depict dominant long. cross-meander flow paths derIved from trackmg in MODFLOW and MODPATH simulations environmental differences can influence the composition, Hyporheic exchange creates environmental patches abundance, and functional attributes of microbial assem­ on the streambed at., I; blages (Halda-Alija et 200 and review by Findlay An alternative way to evaluate the relative importance of and Sobczak, 2000 and references therein), meiofauna the hyporheic zone in streams would be to focus on the (see review by Hakencamp and Palmer, 2000 and refer­ streambed, rather than stream discharge. All hyporheic ences therein), and macro invertebrates (Williams, 1984; exchange (both upwelling and downwelling) must flow Stanford and Ward, 1988). Also, exchange flows of through the benthic zone, the biologically active layer stream water create the environmental conditions nec­ composed of the streambed and the immediately underly­ essary for the incubation of the eggs of that ing sediment. Benthic organisms are directly influenced in the streambed and create the conditions supporting the by hyporheic exchange flows, and in upwelling zones, early life stages of many that reside in streambed upwelling water passes through the benthic zone before gravels (Vaux, 1962; Baxter and Hauer, 2000; Malcolm it is mixed into the . Consequently, simple et at., 2005). budgetary estimates of mass transport may not accu­ The amount of available hyporheic habitat and the rately reflect the relative importance of the hyporheic environmental conditions found there is likely to be deter­ zone to stream ecosystem processes when considering mined by the amount, spatial extent, and of the influence of the hyporheic zone on water temperature hyporheic exchange flows. The hyporheic zone is likely or nutrient cycling. to be restricted to a shallow layer immediately adjacent Hyporheic exchange flows do not occur uniformly over to the wetted stream channel in low gradient rivers with the streambed. Rather, different 'patches' on the stream fine-textured sediment as shown by Duff et at., (1998) bed experience different 'upwelling' or 'downwelling' for the Shingobee River. Conversely, in high-gradient environments (Figure 2; Wondzell et at., 2009b). Up­ rivers with coarse-textured sediment, the hyporheic zone welling environments are of special interest, because may extend more than 1000 m from the wetted chan­ upwelling water has the potential to be thermally or nel as shown by Poole et at. (2008) for the Umatilla chemically distinct from stream water. For example, River. The physiochemical environmental patterns found Arrigoni et ai., (2008) showed that hyporheic exchange within the hyporheic zone are more likely to be a func­ creates a diversity of thermal environments in the main, tion of residence time than flow-path length (Zarnetske side, and spring channels of a large, unconstrained allu­ et at., 2011). While much remains unknown about the vial river during summer baseflow. Ebersole et at. (2003) response of organisms to environmental patterns in the demonstrated that hyporheic upwelling zones provided hyporheic zone, it is clear that the hyporheic zone is a thermal refugia for cold-water fishes in streams on unique and important habitat for a variety of organisms late summer days when discharge was low and ambi­ in many river systems, spanning most, if not all, of the ent stream temperature was high. Valett et at. (1994) state-space depicted in Figure I. showed that hyporheic upwelling zones were enriched

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with nitrate, had higher algal , and after floods, manuscript. This work was supported by the National Sci­ algal biomass recovered more quickly in upwelling zones ence Foundation's Hydrologic Sciences Program (EAR- than in downwelling zones. These results suggest that, 9506669 and EAR-9909564). Data were provided by even where the proportion of stream water exchanged the HJ Andrews Experimental Forest research program, through the hyporheic zone is too small to measurably funded by the National Science Foundation's Long-Term change water temperatures or nutrient concentrations of Ecological Research Program (DEB 08-23380), US For­ the whole stream, hyporheic exchange can create environ­ est Service Pacific Northwest Research Station, and Ore­ mental patches critical to structuring stream ecosystems. gon State University.

Conclusions References Are the effects of the hyporheic zone so highly variable in place and time (or among streams) that a consistent rela­ Arrigoni AS. Poole GC. Mertes LAK. O'Daniel SJ. Woessner WW, Thomas SA. 2008. Buffered, lagged, or cooled? Disentangling hyporheic tionship should not be expected? The conceptual model influences on temperature cycles in stream channels. of the hyporheic zone presented here (Figure 1) suggests Research 44: W09418, DOI:IO.1029/2007WR006480.

that the role of the hyporheic zone in stream ecosys­ Baxter CV, Hauer FR. 2000. Geomorphology, hyporheic exchange, and tems is reasonably predictable. This role does change selection of spawning habitat by bull trout (Salvelinus confluentus). Canadian Journal of Fisheries and Aquatic Sciences 57: 1470-148l. with both location within the stream network as well as with seasonal or storm-driven changes in discharge, but Bianchin M, Smith L, Beckie R. 2010. Quantifying hyporheic exchange in a tidal river using temperature time series. Water Resources Research these dynamics are also reasonably predictable. There can 46: W07507, DOI:IO.102912009WR008365.

be a high degree of reach-to-reach variability created by Cardenas MB. 2009. A model for lateral hyporheic flow based on valley underlying variation in channel morphology that cannot slope and channel sinuosity. Water Resources Research 45: W01501, be predicted from the simple relations shown in Figure 1. DOl:10.1 02912008WR007442. However, the hydraulic and geomorphologic factors driv­ Cardenas MB, Wilson JL, Zlotnik VA. 2004. Impact of heterogeneity, bed forms, and stream curvature on subchannel hyporheic exchange. ing hyporheic exchange has seen considerable research in Water Resources Research 40: DOI:IO.102912004WR003008. recent years (for example, Harvey and Bencala, 1993; Domenico PA, Schwartz FW. 1990. PhYSical and Chemical Hydrogeol­ Kasahara and Wondzell, 2003; Cardenas et al., 2004; ogy. John Wiley and Sons: NY; p. 824.

Wondzell et al., 2009b) so that these controlling factors Duff JH, Murphy F, Fuller CC, Triska FJ, Harvey JW, Jackman AP. are reasonably well understood and are already being 1998. A mini drivepoint sampler for measuring pore water solute concentrations in the hyporheic zone of sand-bottom streams. incorporated in models to estimate hyporheic exchange and Oceanography 43: 1378-1383. flows (Cardenas, 2009). Ebersole JL, Liss WJ, Frissell CA. 2003. Thermal heterogeneity. stream Is the hyporheic zone less important in stream ecosys­ channel morphology, and salmonid abundance in northeastern Ore­ tems than is commonly expected? If importance is sim­ gon streams. Canadian Journal of Fisheries and Aquatic Sciences 60: ply measured as the proportion of stream discharge 1266-1280. exchanged through the hyporheic zone in a given stream Elliott AH, Brooks NH. 1997. Transfer of non-sorbing solutes to a streambed with bed forms: Laboratory experiments. Water Resources reach, then the hyporheic zone will only play a signif­ Research 33: 137-151.

icant role in stream ecosystem processes under limited Findlay S. 1995. Importance of surface-subsurface exchange in stream circumstances, namely, in small streams when discharge ecosystems: The hyporheic zone. Limnology and Oceanography 40: is low. But even then it could be argued that the hyporheic 159-164. zone will have a significant effect on whole network Findlay S, Sobczak WV. 2000. Chapter 12: Microbial Communities in Hyporheic Sediments. pgs. 287-306. In Streams and Ground Waters responses because small streams make up most of the Jones JA, Mulholland PJ (eds). Academic Press: San Diego, CA; p. 425. stream network and most of the watershed drains into Grimm NB, Fisher SG. 1984. Exchange between interstitial and surface these small streams. A broader view of the hyporheic water: Implications for stream metabolism and nutrient cycling. Hydro­ zone would include the diverse array of effects that bioiogia Ill: 219-228. hyporheic exchange flows can support in any given Hakenkamp CC, Palmer MA. 2000. Chapter 13: The Ecology of stream reach-creating unique habitats for a wide diver­ Hyporheic Meiofauna. pgs. 307- 336. In Streams and Ground Waters Jones JA. Mulholland PJ (eds). Academic Press: San Diego, CA; p. 425. sity of organisms either directly in the hyporheic zone or Halda-Alija L, Hendricks SP, Johnston TC. 2001. Spatial and Temporal by modifying the environmental character of upwelling Variation of Enterobacter Genotypes in Sediments and the Underlying and downwelling patches on the streambed. Either way, Hyporheic Zone of an Agricultural Stream. 42: the data, analyses, and conceptual model of the hyporheic 286-294. zone presented here supports the expectation that the Hall RO Jr, Bernhardt ES, Likens GE. 2002. Relating nutrient uptake with transient storage in forested mountain streams. Limnology and hyporheic zone is important to stream ecosystems. Oceanography 47: 255-265.

Harvey JW, Bencala KE. 1993. The effect of streambed topography Acknowledgements on surface-subsurface water exchange in mountain catchments. Water Resources Research 29: 89-98. Special thanks to K. Bencala, M. Gooseff, and B. Jones JB Jr, Holmes RM. 1996. Surface-subsurface interactions in stream McGlynn for helpful discussions and reviews of this ecosystems. Trends in Ecology and Evolution 11: 239-242.

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