Modification of food webs is a frequent cause of shifts in ecosystem states that resist reversal when the food web is restored to its original condition. We used the restoration of the large carnivore guild including gray wolves (Canis lupis), cougars (Felis concolor), and grizzly bears (Ursus arctos horribilis) to the northern range of Yellowstone National Park as a model system to understand how ecosystems might resist reconfiguration after the restoration of apex predators to the food web. The absence of wolves, cougars, and grizzly bears for nearly a century from the northern range was the primary cause of dramatic changes in riparian plant communities. Willows (Salix spp.) were suppressed in height by intense browsing by the dominant herbivore, elk (Cervus canadensis). The loss of activity by beavers (Castor canadensis) coincided with the loss of tall willows. We hypothesized that intense elk browsing interrupted the mutualism between willow and beavers: ecosystem engineering by beavers was a critical component of willow habitat and tall willows were a critical component of habitat for beavers. This interruption made riparian communities resilient to the disturbance caused by the restoration of apex predators. We hypothesized further that reductions in elk browsing attributable to reductions in elk population size were not sufficient to prevent the suppression of willow growth. To test these hypotheses, we conducted a 20-year, factorial experiment that crossed simulated beaver dams with the exclusion of browsing. We found that willows grew to heights expected for restored communities only in the presence of dams and reduced browsing. Willows experiencing ambient conditions remained well below this expectation. We found no difference in heights or growth rates of willows in experimental controls and willows in 21 randomly chosen sites, confirming that the results of the experiment were representative of range-wide conditions. A reorganized community of large herbivores was implicated in the suppression of willow growth. We conclude that the restoration of large carnivores to the food web failed to restore riparian plant communities on Yellowstone's northern range, supporting the hypothesis that this ecosystem is in an alternative stable state caused primarily by the extirpation of apex predators during the early 20th century.

•Soil pitting is an ancient technique for concentrating soil moisture to enable plant establishment and promote plant growth. It is especially effective in arid areas where plant establishment is limited by water availability.•Pits created by digging and mounding action have been shown to be effective. Small pits made by soil compression are not very durable. Larger pits last longer and store more moisture.•In mesic areas or years with above-average soil moisture, pitting may not be needed for plant establishment, and better results may be obtained by drill seeding.•Pitting can help control some non-native annual plants by hindering their seed dispersal and concentrating their seeds within pits. This increases intraspecific competition and limits seed production. The pitting technique has been demonstrated in several studies as an effective control tactic for cheatgrass.•Pitting is a strategy that may become more important as climates become hotter and more variable, as it allows soils to stay wetter for longer periods of time.•Many custom-built machines have been developed to create soil pits efficiently. Commercially available machines are less common. Currently, pitters that dig and mound soil and are suitable for rangelands are not commercially available in North America.

AbstractLand managers in western North America often reverse succession by removing pinyon (Pinus spp.) and juniper (Juniperus spp.) trees to reduce fire risk and increase forage for wildlife and livestock. Because prescribed fire carries inherent risks, mechanical methods such as chaining, roller‐chopping, and mastication are often used. Mechanical methods differ in cost and the size of woody debris produced, and may differentially impact plant and animal responses. We implemented a randomized, complete‐block, split‐plot experiment in December 2011 in the Piceance Basin, northwestern Colorado, USA, to compare mechanical methods and to explore seeding (subplot) interactions. We assessed vegetation 1‐, 2‐, 5‐, and 6‐years post‐treatment, and mule deer (Odocoileus hemionus) response via GPS locations 3–8 years post‐treatment. By 2016, treated plots had 3–5 times higher perennial grass cover and ~10 times higher cheatgrass (Bromus tectorum) cover than untreated control plots. Roller‐chopped plots had both the highest non‐native annual forb cover, and when seeded, the highest density of bitterbrush (Purshia tridentata), a nutritious shrub used by mule deer. Masticated plots had higher bitterbrush use during summer and fall, leaving less forage available for winter. Days of winter mule deer use from GPS point locations in chained and roller‐chopped plots was ~70% higher than in control plots, while winter use in masticated plots was similar to control plots. Mule deer use appears related to a combination of hiding cover, resulting from residual woody debris, and winter forage availability. Roller‐chopped plots provide the best combination of hiding cover and winter forage, but mastication or chaining, applied leaving dispersed security cover, may be better options at large scales or when invasive species concerns exist.

Numerous functional traits have been identified as key contributors to plant performance under drought. However, many of these traits, specifically root traits, are rarely considered in the development of native plant cultivars. In this study, we assessed whether cultivars of the perennial grass Elymus trachycaulus (Slender wheatgrass) developed for drought differ in (a) drought resistance (i.e. a plant's ability to maintain aboveground biomass productivity under water deficit), (b) aboveground and belowground traits, and (c) trait responses to drought from other accessions (i.e. other cultivars, wild accessions). We also evaluated trait plasticity, assessed whether multivariate trait relationships varied between control and drought conditions, and determined which suites of traits are related to drought resistance. We worked with seedlings at two developmental stages to assess whether patterns vary ontogenetically. E. trachycaulus cultivars developed for drought did not differ from other accessions in drought resistance or traits related to drought‐coping strategies. The effects of drought and accession on drought resistance, traits, and trait plasticity varied by developmental stage, but relationships among traits varied little between the two developmental stages. A primary axis of functional variation related to resource acquisition (plant height, root length, root tips) was consistently associated with drought resistance. However, which secondary axes were related to drought resistance varied by developmental stage and moisture condition. Our results suggest that traits and performance of commonly used cultivars ought to be reexamined to determine whether they are actually the best candidates for revegetation projects in specific contexts.

Aims Although leaf and root traits may change considerably throughout plant development, ontogenetic variation is rarely considered in trait-based ecology. Studies focused on how morphological root traits change throughout ontogeny are especially rare. Our objectives were to determine how ontogeny influences seedling traits to inform trait selection for future studies and to advance understanding of how traits at early developmental stages influence seedling growth. Methods We measured traits from eleven herbaceous species at several developmental stages. We used Bayesian random effects models to assess the effects and variation resulting from species identity and ontogeny for each trait. We used principal component analysis and multiple regression to identify which dominant axes of variation were correlated with future growth rates. Results Variation in traits resulting from ontogeny was greatest for growth rates and root elongation rates. Relationships among traits were similar at all ontogenetic stages, but which principal component axes were correlated with future growth depended on stage; at the earliest harvest, the axis related to tissue construction was linked to future growth rate, whereas, at the last harvest, three independent axes were related to future growth rate. Conclusions In our study, traits including leaf dry matter content, root tissue density, and root diameter varied little throughout seedling development and thus may be promising candidates for future trait-based studies. Linking suites of traits to growth strategies may be particularly fruitful for understanding plant strategies throughout early development, as multivariate relationships among traits appear to be more ontogenetically stable than individual traits.

Abstract Understanding the combined effects of drought and invasive species on plant community development and soil moisture could provide valuable insight into the mechanisms hindering successful native plant establishment in dryland restoration projects. We implemented a re‐vegetation experiment at two sites in Colorado, USA (one each in the Western Great Plains and Cold Desert ecoregions) to investigate the effects of drought (66% reduction of ambient growing season rainfall), non‐native Bromus tectorum seed addition (465 seeds/m2), and superabsorbent polymer soil amendment (25 g/m2) on plant community development and soil volumetric water content at 5 and 30 cm depth. Drought resulted in higher B. tectorum cover at the Western Great Plains site but lower B. tectorum cover at the Cold Desert site. These contrasting results suggest drought may interact with site‐specific precipitation patterns to influence B. tectorum establishment. At the Western Great Plains site, drought reduced seeded forb cover and B. tectorum seed addition reduced seeded grass cover, highlighting how the effects of drought and invasive species may vary depending on which functional group is assessed. At the Cold Desert site, drought and B. tectorum seed addition each decreased seeded species cover from approximately 8% to 3%. Superabsorbent polymer effects were limited to slight increases in overall seeded grass cover at the Western Great Plains site from 2.2% to 4.9%. Both drought and B. tectorum seed addition reduced soil volumetric water content but in some cases effects depended on interactions between the two treatments, site, or soil depth. Notably, at the Cold Desert site the reduction in soil volumetric water content resulting from B. tectorum seed addition with ambient precipitation exceeded that of the drought treatment alone at 5 and 30 cm depth. Our results demonstrate that drought and invasive species both negatively influence native plant establishment and soil moisture, and in some cases may interact. Considering both abiotic and biotic stressors as well as their interactions in restoration planning could improve our understanding of native plant establishment and improve re‐vegetation outcomes.

Effective manipulations to prevent the spread of invasive species are needed. Downy brome (Bromus tectorum L.) is an annual invader that often expands after disturbances, compromising restoration of big sagebrush (Artemisia tridentata Nutt.) communities in western North America. This study examined the effects of two manipulations that may slow seed dispersal: soil microtopography (roughened with 50-cm relief or flat) and woody debris (0.024 m3·m− 2 or none) on restoration of four disturbed mountain big sagebrush (A. tridentata Nutt. ssp. vaseyana) sites in Colorado. Treatments were crossed with seeding in a fully factorial experiment (n = 3). Microtopography and woody debris treatments were also crossed in a seed dispersal experiment using fluorescently marked downy brome seeds. In the restoration study, downy brome invaded two sites, one pervasively and one patchily. Seeding limited downy brome cover at both of these sites and also increased perennial grass and forb cover while limiting shrub cover. At the pervasively invaded site, the rough surface reduced unseeded plot downy brome cover from 13% to 3% by 5 yr post treatment. Woody debris increased shrub and perennial grass cover but had little effect on downy brome. In the seed dispersal experiment, the rough surface reduced downy brome mean dispersal distance twofold to threefold and 95% quantile distance threefold to sixfold. Woody debris slightly reduced downy brome dispersal only within rough surface plots. A rough surface may aid restoration by trapping downy brome seeds near the parent plant, limiting their spatial distribution, increasing intraspecific competition, and reducing propagule pressure. Designing landscapes to slow seed dispersal may help control invasives and promote establishment of seeded species.

Water‐holding soil amendments such as super‐absorbent polymer (SAP) may improve native species establishment in restoration but may also interact with precipitation or invasive species such as Bromus tectorum L. (cheatgrass or downy brome) to influence revegetation outcomes. We implemented an experiment at two sites in Colorado, U.S.A., in which we investigated the interactions of drought (66% reduction of ambient rainfall), B. tectorum seed addition (BRTE, 465 seeds/m2), and SAP soil amendment (25 g/m2) on initial plant establishment and 3‐year aboveground and belowground biomass and allocation. At one site, SAP resulted in higher native seeded species establishment but only with ambient precipitation. However, by the third year, we detected no SAP effects on native seeded species biomass. Treatments interacted to influence aboveground and belowground biomass and allocation differently. At one site, a SAP × precipitation interaction resulted in lower belowground biomass in plots with SAP and drought (61.7 ± 7.3 g/m2) than plots with drought alone (91.6 ± 18.1 g/m2). At the other site, a SAP × BRTE interaction resulted in higher belowground biomass in plots with SAP and BRTE (56.6 ± 11.2 g/m2) than BRTE alone (35.0 ± 3.7 g/m2). These patterns were not reflected in aboveground biomass. SAP should be used with caution in aridland restoration because initial positive effects may not translate to long‐term benefits, SAP may uniquely influence aboveground versus belowground biomass, and SAP can interact with environmental variables to impact developing plant communities in positive and negative ways.

Restoring native plants in rangelands threatened by downy brome (Bromus tectorum L.) presents a serious challenge to land managers. Higher, more consistent soil moisture, as well as slightly compacted soils, may reduce the competitive abilities of downy brome. We manipulated these factors with three treatments: superabsorbent polymer (SAP), a soil-binding agent, and roller compaction at two restoration sites, Wagon Road Ridge (WRR) and Sagebrush (SGE), in northwestern Colorado. SAPs absorb water when soils are wet and then gradually release it, often reducing plant water stress. The binding agent we used is purported to increase water infiltration while reducing soil movement. In Experiment 1, we crossed an SAP, a binding agent, and rolling and found that SAP benefitted perennial grass establishment at the WRR site only. SAP also decreased downy brome cover and biomass at WRR. The binding agent increased soil moisture at both sites, and the highest level of binding agent reduced downy brome cover in the absence of SAP at the SGE site. In Experiment 2, we examined only SAP, with larger plots and a more complex seed mix. Again, SAP benefitted perennial grass establishment at WRR only. SAP reduced initial perennial forb density at both sites but did not affect forb cover in subsequent years. SAP effects on downy brome were site-specific. There was a trend for reduced downy brome cover with SAP at WRR, but SAP caused a large increase in downy brome cover in yr 3 at SGE. Granulated SAP can be applied easily along with drill seeding, making it potentially applicable for dryland restoration. However, site specific factors may influence whether perennial grasses or downy brome most benefit from SAP application.

Abstract The generally negative effect of anthropogenic disturbance on the quality of habitats for species viability makes it a common focus of conservation assessment and prioritization efforts. Although many available spatial models and metrics (e.g., distance to or density of disturbance) characterize impact patterns of anthropogenic disturbance on the landscape, a general evaluation of model performance against empirical measurements of ecological integrity is lacking. We tested both distance‐based and disturbance‐density models in relation to ecological indicators. The models included roads, residential and commercial development, agricultural land use, mining, energy development infrastructure, and transmission structures as disturbance sources. Model parameters were based on expert input and results from the published literature. The disturbance models were tested against two disparate and independent measures of habitat quality: a floristic quality index and measures of greater sage‐grouse population integrity. Floristic quality scores were significantly lower in vegetation plots closer to disturbances in a general distance‐based disturbance model across Colorado. Although the proportion of variation in floristic quality explained by anthropogenic disturbance was relatively low (8.5–11.8%), it appeared to represent a ubiquitous baseline negative effect of proximity to anthropogenic disturbance on the quality of vegetation communities. For both distance‐ and density‐based greater sage‐grouse models, modeled disturbance indices were significantly lower (10–12 times) near active than historic leks, and numbers of males counted at leks increased significantly (3.2–3.4 times) as modeled disturbance decreased. Our findings indicate that as a general class, geospatial models can depict effects of anthropogenic disturbance on both plant communities and individual animal species. Empirical validation of disturbance models focused on other species or regions is recommended to further evaluate the utility and reliability of these methods.

Pinyon-juniper (Pinus spp.–Juniperus spp.) encroachment and declining mule deer (Odocoileus hemionus) populations in western Colorado have necessitated management for increased forage. Pinyon-juniper removal is one such technique; however, it is unclear which method of tree removal most effectively promotes forage species. We conducted an experiment to quantify understory responses to mechanical pinyon-juniper removal and seed additions in a blocked design using three different methods: anchor-chaining, rollerchopping, and mastication. Blocks contained each mechanical and seeding treatment along with an untreated control. Seven blocks across two sites, North Magnolia (NM, 4 blocks) and South Magnolia (SM, 3 blocks), were treated during the fall of 2011. Half of each plot was seeded before or during mechanical treatment with a mix of grasses, shrubs, and forbs. After two growing seasons, biomass of perennial grasses was 90–160 kg · ha−1 in mechanically treated plots compared with 10 kg · ha−1 in untreated controls. There were no differences, however, between mechanical treatments for any perennial plant species. Response of annual plant species depended on mechanical treatment type and site. Rollerchopping had higher exotic annual grass cover than mastication or control at NM and higher exotic annual forb cover than chaining or control at SM. Rollerchopping was the only treatment to have higher native annual forb cover than control in the absence of seeding. Seeding increased native annual forb biomass in mastication compared with control. Seeding also increased shrub density at SM, which had fewer shrubs pretreatment relative to NM. Results suggest any type of mechanical removal of pinyon-juniper can increase understory plant biomass and cover. Seeding in conjunction with mechanical treatments, particularly mastication, can initially increase annual forb biomass and shrub density. Finally, different understory responses between sites suggests that pretreatment conditions are important for determining outcomes of pinyon-juniper removal treatments.

Energy-extraction disturbances entail soil handling and often have large edge-to-area ratios. These characteristics should be considered when designing weed-control strategies. In western North America, many energy developments coincide with infestations of downy brome, an annual grass that severely curtails productivity, diversity, and habitat value of invaded areas. Downy brome is sensitive to soil compaction and seed burial, both of which may occur when soil is handled. In this study, I examined the effect of soil-density manipulations and herbicide application (105 g ai ha −1 imazapic with 280 g ai ha −1 glyphosate) on six simulated pipeline disturbances in a Wyoming big sagebrush ecosystem invaded by downy brome. Disturbances occurred at the end of the growing season, after ambient downy brome seed rain in the study areas had abated. Treatments and seeding occurred shortly after disturbances. The following spring, downy brome seedling density was 10-fold lower within disturbances than in control areas, but seedling density quickly rebounded in disturbed areas where no herbicide had been applied. In herbicide plots, downy brome seedling density remained low during the first growing season, and shrub cover after 3 yr was eight times higher than in no-herbicide plots. Soil density manipulations via disking and rolling treatments had little effect on downy brome. Prior research has shown that imazapic is more effective when combined with disturbances, such as fire. This study demonstrates that imazapic may also be effective in combination with a disturbance that is timed to bury downy brome seeds.

Abstract Because of disturbance and exotic plant invasions, ecological restoration is necessary for maintaining functional big sagebrush ecosystems in western North America. Downy brome control is often necessary in restoring this ecosystem type; however, many brome control measures hinder ecological restoration by limiting the types of plants which can be established. Microtopography manipulation may aid weed control by entrapping undesirable seeds. We undertook a field experiment at four sites in the Piceance Basin of western Colorado, USA to test the effects of microtopography (rough with brush mulch or flat with straw mulch), seed mix (high-forb or balanced), and herbicide (140 g ai ha−1 imazapic ammonium salt or none) on downy brome control and perennial plant establishment following disturbance. Three years post-treatment, downy brome had become established at two of the four sites, one each with high (GVM) and low (MTN) downy brome seed rain. At GVM, the rough/brush treatment augmented the effectiveness of imazapic, reducing downy brome biomass six-fold. At MTN, the rough/brush surface reduced downy brome biomass 10-fold in the absence of imazapic. Across all four sites, forb and shrub biomass were higher with the high-forb mix, and there was no effect of seed mix on downy brome or annual forb biomass. Restoring a full complement of plant functional groups in big sagebrush ecosystems may be aided by increasing forbs in seed mixes, and manipulating soil microtopography. Nomenclature: imazapic ammonium salt, big sagebrush, Artemisia tridentata Nutt. ARTR2, downy brome, Bromus tectorum L. BROTE. Management Implications: Big sagebrush ecosystems in western North America are threatened by many factors, including downy brome invasion, conifer encroachment, and disturbance. Restoring downy brome-invaded ecosystems is difficult because many effective downy brome control measures, such as seeding competitive grasses or applying herbicides, impair the growth of desirable forbs and shrubs. Previous work has shown that downy brome seeds disperse more readily when the soil surface is flat or when vegetation is removed, such as following a fire or after the creation of an oil and gas well pad. In this study, we examined whether creating obstructions to seed dispersal could improve restoration. We tested three factors: (1) a rough soil surface, comprised of pothole-sized holes and mounds with brush mulch vs. a flat soil surface with straw mulch, (2) a high-forb seed mix, containing nearly 75% forbs by seed number, vs. a mix containing roughly equal numbers of forb, grass, and shrub seeds, and (3) imazapic herbicide at 140 g ai ha−1 vs. no herbicide. Downy brome established at two of the four sites, and at both of these, the rough surface helped to control downy brome. The rough surface combined with imazapic was most effective at a site that had some downy brome prior to disturbance, while the rough surface alone was effective at another site which was not invaded prior to disturbance. It is possible that the rough surface makes downy brome less competitive because the seeds get trapped in the holes, which have higher soil moisture, as downy brome is typically less competitive in wetter environments. The high-forb seed mix resulted in higher forb and shrub establishment, and lower grass establishment, than the balanced mix. Even so, grass cover was still higher than forb cover at most sites, and was higher than in the undisturbed communities. There was no difference in downy brome or weedy annual forbs due to seed mix, and the high forb cover produced by the high-forb mix is beneficial for sage-grouse habitat restoration. Seeding mostly forbs and shrubs at a high rate, 1600 seeds m−2, should be considered in areas where erosion is not a concern. The imazapic treatment did control downy brome, but it also had a negative impact on grasses, forbs, and shrubs. The lack of perennial plant establishment led to higher annual forb cover in imazapic plots three years post-treatment. Extending the plant-back interval, applying at a lower rate, and restricting application to areas with downy brome presence prior to disturbance are recommended.

Particularly well-known among the many impacts of the invasive annual grass downy brome (Bromus tectorum, Poaceae) is its ability to alter fire cycles and increase in abundance after fire. However, little is known about how fire influences B. tectorum dispersal. We quantified fire effects on B. tectorum dispersal using three recently burned areas in the western region of the Colorado Rocky Mountains by marking diaspores (seeds) with fluorescent powder, and then recovering them at night using ultraviolet lights. Diaspores were of two types: with and without sterile florets attached. We also characterized vegetation cover and near-surface wind speed in burned and unburned areas. Diaspores travelled much farther in burned areas than in nearby unburned areas (mean ± standard error at the end of the experiment: 209 ± 16 cm and 38 ± 1 cm, respectively; maximal distance at the end of the experiment: 2,274 cm and 150 cm, respectively), indicating an increase in dispersal distance after fire. Diaspores with sterile florets attached dispersed longer distances than those without sterile florets (mean ± standard error at the end of the experiment: 141 ± 14 cm and 88 ± 7 cm, respectively). Vegetation cover was lower and wind speeds were higher in the burned areas. Our results indicate that at least one of the mechanisms by which the spread of B. tectorum is promoted by fire is through increased seed dispersal distance. Preventing movement of seeds from nearby infestations into burned areas may help avoid the rapid population expansion often observed.

The presence or absence of obstructions can affect seed dispersal. Reclamation activities often cause changes in the type and amount of such obstructions. The consequences of removing obstructions on the dispersal of undesirable species are unknown. In western North America, reclamation may often proceed in areas surrounded by the invasive Bromus tectorum L. (cheatgrass). The importance of preventing cheatgrass seed dispersal from surrounding landscapes is an unknown factor in reducing cheatgrass competition in these areas. I quantified cheatgrass seed dispersal in the early stages of reclamation, when soils were bare. Four groups of 100 sterilized, fluorescently marked cheatgrass seeds were released in each of three areas in NW Colorado, USA. Seeds were located at night using blacklights four times over 14 days, and the distance between each seed and the point of release was measured. Across sites, dispersal distance averaged 2.4 m, 5% of seeds traveled further than 7.6 m, and maximum recorded distance was 20.8 m. Maximum distances reported in this study are 50‐fold higher than previously reported for intact sagebrush ecosystems. I suggest that the difference is due to a lack of impediments to secondary dispersal on bare soil. When reclamation areas are surrounded by weeds such as cheatgrass, seeds dispersing from the perimeter may influence restoration outcome.

Excessive levels of herbivory, incision of stream channels, and climate warming are believed to be responsible for the decline of woody deciduous plants in riparian zones in western North America, declines that are likely to be associated with diminished biological diversity. In the northern elk wintering range of Yellowstone National Park, USA, overbrowsing by elk (Cervus elaphus), lowered water tables resulting from stream incision, and loss of activity by beaver (Castor canadensis) have been implicated in the decline of willow (Salix sp.) communities. Reducing elk browsing appears sufficient for willow recovery in some areas, but where water table changes have been dramatic, recovery may be slow or absent. The importance of water table changes is disputed because experimental results demonstrate water table limitations, but water table depth has failed to explain variation in willow height at landscape scales. One explanation for this apparent discrepancy is that willows that have survived intensive browsing by elk have maintained access to groundwater despite declining water tables. Using stable isotopes of water, we examined the relationships between groundwater use, water table depth, and height of heavily browsed Salix geyeriana. Salix geyeriana groundwater use varied from 30% to 80%, and was higher later in the growing season, when soil water was less available and shoot water potentials were lower. Late season groundwater use explained 26% of the variation in total height of willows (P = 0.002), with taller plants using more groundwater. Water table depth explained only 8% of the variability in total height (P = 0.051), with shorter willows having deeper water table depths. Groundwater use and water table depth were uncorrelated. Height recovery following a winter of heavy browsing was related to groundwater use, but not groundwater depth. We suggest that access to deeper water sources alleviates late season water stress, allowing for more rapid height recovery and higher total plant height. Variability in groundwater access may account for variability in height recovery at landscape scales.

In the northern elk wintering range of Yellowstone National Park, USA, wolf (Canis lupus) removal allowed elk (Cervus elaphus) to overbrowse riparian woody plants, leading to the exclusion of beaver (Castor canadensis) and a subsequent water table decline in many small stream valleys. Reduced elk browsing following wolf reintroduction may or may not facilitate willow (Salix sp.) recovery in these areas. To determine if the effect of elk browsing on willow interacts with that of beaver abandonment, we manipulated elk browsing and the water table in a factorial experiment. Under the condition of an ambient (low) water table, elk browsing increased shoot water potential (Psis), photosynthesis per unit leaf area (A), stomatal conductance per unit leaf area (gs), and aboveground current annual growth (CAG) by 50%. Elk browsing occurred entirely during dormancy and did not affect total plant leaf area (L). Improved water balance, photosynthetic rate, and annual aboveground productivity in browsed willows appeared to be due to morphological changes, such as increased shoot diameter and decreased branching, which typically increase plant hydraulic conductivity. An elevated water table increased Psis, A, gs, CAG, and L, and eliminated or lessened the positive effect of browsing on CAG for most species. Because low water tables create conditions whereby high willow productivity depends on the morphological effects of annual elk browsing, removing elk browsing in areas of water table decline is unlikely to result in vigorous willow stands. As large willow standing crops are required by beaver, a positive feedback between water-stressed willow and beaver absence may preclude the reestablishment of historical conditions. In areas with low water table, willow restoration may depend on actions to promote the re-establishment of beaver in addition to reducing elk browsing.