Importance of low-flow and high-flow characteristics to restoration of riparian vegetation along rivers in arid south-western United States

1. Riparian vegetation in dry regions is influenced by low‐flow and high‐flow components of the surface and groundwater flow regimes. The duration of no‐flow periods in the surface stream controls vegetation structure along the low‐flow channel, while depth, magnitude and rate of groundwater decline influence phreatophytic vegetation in the floodplain. Flood flows influence vegetation along channels and floodplains by increasing water availability and by creating ecosystem disturbance. 2. On reference rivers in Arizona's Sonoran Desert region, the combination of perennial stream flows, shallow groundwater in the riparian (stream) aquifer, and frequent flooding results in high plant species diversity and landscape heterogeneity and an abundance of pioneer wetland plant species in the floodplain. Vegetation changes on hydrologically altered river reaches are varied, given the great extent of flow regime changes ranging from stream and aquifer dewatering on reaches affected by stream diversion and groundwater pumping to altered timing, frequency, and magnitude of flood flows on reaches downstream of flow‐regulating dams. 3. As stream flows become more intermittent, diversity and cover of herbaceous species along the low‐flow channel decline. As groundwater deepens, diversity of riparian plant species (particularly perennial species) and landscape patches are reduced and species composition in the floodplain shifts from wetland pioneer trees (Populus, Salix) to more drought‐tolerant shrub species including Tamarix (introduced) and Bebbia. 4. On impounded rivers, changes in flood timing can simplify landscape patch structure and shift species composition from mixed forests composed of Populus and Salix, which have narrow regeneration windows, to the more reproductively opportunistic Tamarix. If flows are not diverted, suppression of flooding can result in increased density of riparian vegetation, leading in some cases to very high abundance of Tamarix patches. Coarsening of sediments in river reaches below dams, associated with sediment retention in reservoirs, contributes to reduced cover and richness of herbaceous vegetation by reducing water and nutrient‐holding capacity of soils. 5. These changes have implications for river restoration. They suggest that patch diversity, riparian plant species diversity, and abundance of flood‐dependent wetland tree species such as Populus and Salix can be increased by restoring fluvial dynamics on flood‐suppressed rivers and by increasing water availability in rivers subject to water diversion or withdrawal. On impounded rivers, restoration of plant species diversity also may hinge on restoration of sediment transport. 6. Determining the causes of vegetation change is critical for determining riparian restoration strategies. Of the many riparian restoration efforts underway in south‐western United States, some focus on re‐establishing hydrogeomorphic processes by restoring appropriate flows of surface water, groundwater and sediment, while many others focus on manipulating vegetation structure by planting trees (e.g. Populus) or removing trees (e.g. Tamarix). The latter approaches, in and of themselves, may not yield desired restoration outcomes if the tree species are indicators, rather than prime causes, of underlying changes in the physical environment.     

Limited Effects of Large‐Scale Riparian Restoration on Seed Banks in Agriculture

I examined the effect of riparian forest restoration on plant abundance and diversity, including weed species, on agricultural lands along the Sacramento River in California (United States). Riparian forest restoration on the Sacramento River is occurring on a large‐scale, with a goal of restoring approximately 80,000 ha over 160 km of the river. In multiuse habitats, such as the Sacramento River, effects of adjoining habitat types and movement of species across these habitats can have important management implications in terms of landscape‐scale patterns of species distributions. Increased numbers of pest animals and weeds on agricultural lands associated with restored habitats could have negative economic impacts, and in turn affect support for restoration of natural areas. In order to determine the distribution and abundance of weeds associated with large‐scale restoration, I collected seed bank soil samples on orchards between 0 and 5.6 km from adjacent restored riparian, remnant riparian, and agricultural habitats. I determined the abundance, species richness, and dispersal mode of plant species in the seed bank and analyzed these variables in terms of adjacent habitat type and age of restored habitat. I found that agricultural weed species had higher densities at the edge of restored riparian habitat and that native plants had higher densities adjacent to remnant riparian habitat. Weed seed abundance increased significantly on walnut farms adjacent to restored habitat with time since restored. I supply strong empirical evidence that large areas of natural and restored habitats do not lead to a greater penetration of weed species into agricultural areas, but rather that weed penetration is both temporally and spatially limited.     

Waste Not,Want Not: The Need to Utilize Existing Artificial Structures for Habitat Improvement Along Urban Rivers

Urban rivers have often experienced substantial engineering modification and consequently are highly degraded aquatic ecosystems with minimal riparian habitat. Habitat restoration and improvement efforts are needed within urban rivers to support ecological communities and increase ecosystem integrity. Most river restoration techniques are not feasible within large urban rivers, and so there is a need to develop novel methodologies. Artificial structures such as river walls can function as habitat for plant and invertebrate species in urban rivers, and in some cases can be more diverse than remnant habitat. Along the River Thames through central London, plant species richness was found to be significantly higher on river walls than intertidal foreshore, which represents the only remnant habitat for riparian species. Both this survey and other studies have suggested that the physical and environmental characteristics of river walls are likely to influence their capacity to function as ecological habitat, for example, walls composed of more complex construction materials (brick and boulders) being more diverse than simpler structures (concrete and sheet piling). The opportunity exists to use river walls and other artificial structures (e.g., jetties) to improve habitat along urban rivers by installing walls which are designed to be more complex, or by adding modifications to existing walls. Some trial modifications, such as the addition of wall ledges and timber fenders to sheet piling, have been installed at Deptford Creek along the River Thames, and have so far greatly supported the colonization and development of plant communities. The restoration possibilities of such modifications should be considered, and further development and rigorous testing of installations is required in urban rivers to make sound restoration recommendations.     

Flood‐deposited wood creates regeneration niches for riparian vegetation on a semi‐arid South African river

Question : The formation of large woody debris (LWD) piles during floods has significant impacts on riparian succession through pioneering plants often establishing in association with wood. We assess the importance of LWD for seed regeneration of riparian plants after a century‐scale flood disturbance in a semi‐arid environment. Location : The Sabie River within Kruger National Park in the semi‐arid northeast of South Africa. Methods : Our approach was to quantify the riparian soil seed bank, to record the frequency of establishment of riparian plants in woody debris piles, and to conduct experimental out‐plantings of common riparian trees in plots with and without LWD. Results : We found the abundance and diversity of seedlings were higher in soils taken from wood piles than from open reference areas, and most seedlings were herbaceous species. Surveys indicated that numbers of seedlings recorded within woody debris were significantly greater than in open reference areas or within established vegetation. Seedling establishment in various cover‐types also varied for different riparian tree species. Experimental out‐planting of seedlings of two riparian tree species (Philenoptera violacea and Combretum erythrophyllum) revealed that, after 433 days, planted seedlings survived only in woody debris piles. Conclusion : LWD formed after a large flood creates heterogeneous patches that may influence post‐disturbance regeneration of riparian vegetation by providing a variety of environmental niches for seedlings establishment. We suspect that higher seedling survival in LWD is due to increased moisture (particularly in the dry season) and nutrients, and protection from seasonal flooding and herbivory.     

Soil seed banks of two montane riparian areas: implications for restoration

Understanding the role of seed banks can be important for designing restoration projects. Using the seedling emergence method, we investigated the soil seed banks of two montane, deciduous riparian forest ecosystems of southeastern Arizona. We contrasted the seed banks and extant vegetation of Ramsey Canyon, which is the site of riparian restoration activities, with that of Garden Canyon, which has been less affected by human land uses. Fewer plant species were found at Ramsey Canyon than Garden Canyon, for both the seed bank and extant vegetation, and the vegetation at Ramsey Canyon (seed bank and extant) had consistently drier wetland indicator scores. As well, vegetation patterns within sampling zones (channel margins and adjacent riparian forests) differed between canyons. At Garden Canyon channel margins, the seed bank and extant vegetation had relatively high similarity, with herbaceous wetland perennial species dominating. Extant vegetation in the floodplain riparian forest zone at Garden Canyon had a drier wetland indicator score than the seed bank, suggesting that the floodplains are storing seeds dispersed from wetter fluvial surfaces. Vegetation patterns for Ramsey Canyon channel margins were similar to those for Garden Canyon floodplains. Vegetation patterns in the Ramsey Canyon riparian forest zone were indicative of non-flooded conditions with an abundance of upland species in the soil seed bank and extant vegetation. Channel geomorphology measurements indicated that much of the riparian forest zone at Ramsey Canyon is functionally a terrace, a condition that may be a legacy of channel erosion from historic land uses. Steep, erodible channel slopes may contribute to the low seed bank germinant density at Ramsey Canyon channel margins, and narrower flood-prone area may explain the greater terrestrialization of the vegetation in both sampling zones. We recommend testing the use of donor soils from more diverse stream reaches to restore biodiversity levels at Ramsey Canyon, following restoration activities such as channel-widening. Seed banks from Garden Canyon, for example, although predominantly consisting of herbaceous perennials, would supply species with a range of moisture tolerances, life spans, and growth forms. We also recommend that restorationists take care not to harm seed banks exposed during removal of introduced species; at Ramsey Canyon, soil seed banks were equally diverse in areas with high and low cover of the introduced Vinca major (a legacy of Ramsey Canyon land use).     

Seed bank,seed dispersal and vegetation cover: Colonization along a newly‐created river channel

Question : What is the relative importance of the initial seed bank and subsequent seed dispersal for floristic composition of bank vegetation two years after creation of a newly‐cut reach of a river channel? Location : River Cole, West Midlands, United Kingdom. Methods : We took bank and bed sediment samples from a 0.5‐km reach of a new river channel cut into intact flood‐plain. After river diversion, seed samples deposited on artificial turf mats placed on the river banks and flood‐plain edge were taken in summer and winter 2002 and 2003. Seed rain samples from funnel traps were taken during summer 2002 and 2003. We undertook greenhouse germination trials to assess viable seed species within these samples. In summer 2004, we surveyed river bank vegetation. Agglomerative cluster analysis was used to investigate floristic similarity between seed bank, seed rain, seed deposition samples and final bank vegetation cover. DCA was used to explore contrasts between the samples and to assess whether these reflected interpretable environmental gradients. Results : Seed rain samples contained a small subset of species in the summer depositional samples. 38 species were found within the final vegetation, the seed bank, and at least one of the four sets of depositional samples; a further 30 species not present in the seed‐bank samples were present in at least one of the four sets of depositional samples and the final vegetation. Floristic composition of the vegetation was most similar to the depositional samples from winter 2002 and 2003 and summer 2003. DCA axis 1 reflected a time sequence from seed‐bank samples through depositional samples to the final vegetation. Conclusions : Newly cut river banks were colonized rapidly. Seed remobilization and hydrochorous transport from the upstream catchment are important for colonization. Species richness was highest in samples deposited during winter when high river flows can remobilize and transport viable seeds from upstream. This process would also have enhanced the species richness of seed production along the banks during the second summer (2003).     

Plant species richness in ephemeral and perennial reaches of a dryland river

Ephemeral reaches are common along desert rivers but are less well studied than those with perennial stream flow. This study contrasted riparian plant species richness and composition (extant vegetation and soil seed bank) between stream reaches with different low-flow conditions (perennial vs. ephemeral flow) but similar flood patterns and similar watershed-derived species pools. Data were collected at Cienega Creek (Arizona, USA) over a 2 year period spanning drought conditions and wetter conditions. Consistent with expectations relating to water limitation effects on diversity, species richness in the riparian zone was lower at ephemeral-flow sites during a season with minimal precipitation and no overbank flooding; under these conditions, the more permanent water sources of the perennial-flow sites sustain the larger number of species. During seasons with greater precipitation and elevated stream flows, in contrast, species richness at ephemeral-flow sites increased to levels at or slightly above those of perennial-flow sites. For values pooled across two wet seasons of a calendar year, year-round richness was greater at the two ephemeral-flow sites (total of 92 vascular plant species) than at the two perennial-flow sites (68 species). This greater year-round richness was a combination of multiple factors: greater light, space, and bare ground, a diverse soil seed bank (with the seed banks equally species-rich among hydrologic types), and moderately abundant precipitation and flooding sufficient to stimulate establishment of opportunistic species (mainly annuals) during the bimodal wet seasons. These results indicate that long-term patterns of site water availability, by influencing woody plant cover, mediate the diversity response to episodic water pulses in dryland rivers. The results also have implications for riparian conservation efforts, which to date have focused primarily on perennial stream reaches: ephemeral reaches of spatially intermittent rivers harbor many riparian plant species, and warrant conservation efforts, as well.     

Soil seed bank in a patchy vegetation of coastal sandy plains in southeastern Brazil

Large seed banks have been found in tropical dry forests and also in habitats with high seasonality in rainfall. However, patchily structured vegetation could induce great spatial variation in the seed bank. We characterized the seed bank in a patchy vegetation of restinga, a common type of coastal vegetation found in the Atlantic forest biome. We also evaluated whether there is any spatial variation between the litter and soil layer, bare sand, and the edge and center of vegetation patches with distinct species dominance. We found 104 seeds/m² in the seed bank using a 5‐cm‐depth sampling. Seven out of 16 species found in the restinga seed bank germinated; two of these were found in the early stages of vegetation patches. We found a higher number of seeds at the edge than in the center of vegetation patches. However, there were no significant differences in the number of seeds in the seed bank between the litter and soil layer, and between vegetation patches with distinct species dominance. Bare sandy soils had lower seed bank densities than vegetation patches. A small seed bank size might be explained by the low proportion of seeds from herbaceous and woody species, which are pioneers in the Atlantic forest. However, seed bank might play an important role in the early stages of the successional process, due to the occurrence of the few species that are able to colonize new young vegetation patches.     

Does the seed bank contribute to the restoration of species‐rich vegetation in wet dune slacks?

Questions: What is the contribution of the seed bank to restoration of species‐rich vegetation in oligotrophic wet dune slacks? Does the restoration management affect the seed bank? Location: Calcareous coastal dune slacks at the west coast of The Netherlands. Methods: Species composition of the seed bank and the above‐ground vegetation was sampled in dune slacks that had a variable extent of groundwater level rise in combination with either topsoil removal or mowing. Results: The seed bank had a high potential for restoration of species‐rich vegetation: 60 species were found in the seed bank of which 14 were characteristic of oligotrophic, wet dune vegetation. While topsoil removal almost completely removed the seed bank, groundwater level rise did not permanently submerge the seed bank of species of oligotrophic, wet conditions. Changes in abundance in the established vegetation were unrelated to species abundance in the seed bank. Of all new species establishments in the vegetation relevés, 76% occurred where the species was not found in the seed bank. The chance that presence of a species in the seed bank led to establishment the following year was only 11%. Conclusion: The seed bank was not the dominant source for newly establishing species following the large disturbance that was induced by restoration management. Changes in species abundance after the restoration impact were not related to species abundance in the seed bank, but to ongoing succession and current year dispersal. To attain a high number of new establishments, restoration projects should preferably be planned in the proximity of refuge populations, rather than relying on the seed bank alone.     

Comparison of Designed Channel Restoration and Riparian Buffer Restoration Effects on Riparian Soils

Stream restoration is often employed in efforts to stabilize eroding channel banks. Banks are stabilized through a designed channel approach, which involves grading and armoring of stream banks using heavy machinery, or alternatively through planting of seedlings and saplings to establish forested riparian buffers. We hypothesized that designed channel restoration would have detrimental impacts on riparian soils but that soils would recover over time, and we hypothesized that riparian buffer restoration would not impact riparian soils. We tested these hypotheses by comparing soil attributes (bulk density, soil organic matter, and root biomass) at reaches that had undergone designed channel and riparian buffer restoration in different years (project ages ranged from 2 to 16 years) to paired urban (unrestored) control reaches. Soil properties in sub‐surface soil layers (10–20 and 20–30 cm depth) at both recent (<10 years old) and older (>10 years old) designed channel reaches differed significantly from paired urban control soils; bulk density was higher and root biomass lower in manipulated reaches compared to urban control reaches. At many designed channel reaches, bulk density exceeded values known to restrict root growth. These results indicate that compaction and disturbance of riparian soils may be a significant unintended consequence of designed channel restoration and can persist for at least a decade. In contrast, we found no significant differences in soil properties between riparian buffer restoration reaches and urban control reaches. Thus, the results indicate that riparian buffer restoration is a more ecologically favorable method than designed channel restoration for bank stabilization.