The Role of Abandoned Channels as Refugia for Sustaining Pioneer Riparian Forest Ecosystems

In disturbance-prone ecosystems, organisms often persist in spatial refugia during stressful periods. A clear example is the colonization of abandoned river channels by pioneer riparian trees. Here, we examine the prominence of this establishment pathway for a foundation tree species (Fremont cottonwood, Populus fremontii) within the riparian corridor of a large river, the Sacramento River in central California. We quantified the total proportion of forest that initiated as a result of channel abandonment for a 160-km reach, analyzed concurrent patterns of tree establishment with floodplain accretion and sedimentation history, and developed a conceptual model of biogeomorphic evolution of abandoned channels. Historical air photo analysis indicated that stands associated with abandoned channels comprised more than 50% of the total extant cottonwood forest area. Tree-ring evidence showed that cottonwood stands commonly developed immediately following abandonment, and the recruitment window ranged from 4 to 40 years, but was less than 10 years at most sites. Rates of floodplain rise and fine sediment accumulation were high in young sites and decreased logarithmically over time. Together, these results suggest that abandoned channels are an important refuge for cottonwood recruitment, that the greatest opportunity for colonization occurs within a short period after the cutoff event, and that sedimentation processes influence the duration of the colonization window. On rivers where tree recruitment along the active channel is severely limited by hydrologic regulation and/or land management, abandoned channel refugia may play an even more important role in sustaining an ecologically functional riparian corridor. Preserving bank erosion, active meander corridors and forest regeneration zones created by cutoff events are therefore key conservation measures on shifting rivers.     

Hydroclimatic drivers of the growth of riparian cottonwoods at the prairie margin: River flows,river regulation and the Pacific Decadal Oscillation

Cottonwoods, riparian poplars, are facultative phreatophytes and can obtain water from shallow soil moisture originating from rainfall, or from the deeper capillary fringe above the alluvial water table that is recharged by river water infiltration. The correspondence between cottonwood growth and river flows should reveal the dependency upon alluvial groundwater and subsequently, the vulnerability to reduced river flows. To explore this association, we analyzed historic growth patterns of plains cottonwoods (Populus deltoides) along the Red Deer River (RDR), which is at the northwestern limit of the North American Great Plains. We developed chronologies of yearly radial increments (RI) and basal area increments (BAI) and explored correspondences with the environmental records from the past century. In this semi-arid region, the RI or BAI were not correlated with local precipitation while negative correlation with growth season temperature (T) (r = −0.37, p < 0.01) could reflect reduced growth with hot summers. There was correlation between growth and annual river discharge (Q, and particularly log Q that approximates river stage) and this increased with two year averaging (r = 0.51, p < 0.01), reflecting carry-over in the watershed hydrology and in the ecophysiological response. There was correspondence with the Pacific Decadal Oscillation index (PDO, r = −0.45, p < 0.01), which provides multi-decade transitions that influence Rocky Mountain headwater precipitation and other weather characteristics, and river flows. The combination of Q, PDO and T provided the strongest multiple regression model, accounting for 44% of the historic growth variation (52% correspondence for 1953–2013). The RDR was dammed in 1983, enabling winter flow augmentation, but summer flows were sustained and cottonwood growth and the streamflow correspondence persisted. This indicates that it is the pattern of dam operation and not damming per se that determines the fate of established riparian cottonwoods downstream. This study revealed that these cottonwoods are phreatophytic and dependent upon alluvial groundwater that is recharged from the river. This provides a research strategy to determine whether riparian woodlands along other regulated rivers are similarly groundwater-dependent and could be vulnerable to river flow reductions from excessive water withdrawal for irrigation or other uses, or with climate change.     

C : N : P stoichiometry of dominant riparian trees and arthropods along the Middle Rio Grande

1. We examined the role of flooding on the leaf nutrient content of riparian trees by comparing the carbon : nitrogen : phosphorus (C : N : P) ratio of leaves and litter of Rio Grande cottonwood (Populus deltoides ssp. wislizenii) in flood and non‐flood sites along the Middle Rio Grande, NM, U.S.A. The leaf C : N : P ratio was also examined for two non‐native trees, saltcedar (Tamarix chinensis) and Russian olive (Elaeagnus angustifolia), and six species of dominant riparian arthropods. 2. Living leaves and leaf litter of cottonwoods at flood sites had a significantly lower leaf N : P ratio and higher %P compared with leaves and litter at non‐flood sites. A non‐flood site downstream from wastewater effluent had a significantly lower litter C : N ratio than all other sites, suggesting N fertilisation through ground water. The non‐native trees, saltcedar and Russian olive, had higher mean leaf N content, N : P ratio, and lower C : N ratio compared with cottonwoods across study sites. 3. Riparian arthropods ranged from 5.2 to 7.1 for C : N ratio, 56–216 for C : P ratio, and 8.9–34 for N : P ratio. C content ranged from 25 to 52% of dry mass, N content from 4.7 to 10.8%, and P content from 0.59 to 1.2%. Differences in stoichiometry between high C : nutrient leaf litter and low C : nutrient invertebrates suggests possible food‐quality constraints for detritivores. 4. These results suggest that spatial and temporal variation in the C : N : P ratio of cottonwood leaves and leaf litter is influenced by surface and subsurface hydrologic connection within the floodplain. Reach‐scale variation in the elemental composition of riparian organic matter inputs may have important implications for decomposition, nutrient cycling, and food webs in river floodplain systems.     

Root architecture of riparian trees: river cut-banks provide natural hydraulic excavation, revealing that cottonwoods are facultative phreatophytes

Plant root architecture reveals the sources of water and nutrients but tree root systems are large and difficult to analyze. With riparian (floodplain) trees, river cut-banks provide natural hydraulic excavation of root systems and this presents a unique study opportunity. Subsequently, we developed the ‘Cut-bank Root Method’, a simple, quantitative approach for analyzing the distribution of coarse roots, based on analyses of photographs of river cut-banks. These reveal the vertical extent of roots and median root depths (R_d). We applied this method along six rivers draining the Canadian Rocky Mountains and observed tenfold difference in R_d. The floodplain forests were dominated by cottonwoods and from mountain to prairie zones we observed progressively deeper roots of Populus trichocarpa (black cottonwood, R_d ~ 0.3 m), P. balsamifera (balsam poplar), P. angustifolia (narrowleaf cottonwood), and P. deltoides (prairie cottonwood, R_d ~ 0.9 m), which had R_d similar to P. fremontii (Fremont cottonwood) in Nevada, USA. Roots were shallower for co-occurring facultative riparian trees, with R_d ~ 0.1 m for P. tremuloides (trembling aspen) and Picea glauca (white spruce). Across the Canadian sites, R_d for cottonwoods were strongly associated with a growth season moisture index (May through September precipitation—potential evapotranspiration; R² = 0.97, P < 0.001). Thus, in wetter climates, riparian cottonwoods were shallow-rooted and would be more dependent upon rain than stream flow. Conversely, in the drier semi-arid regions the cottonwoods were phreatophytic, with deeper root systems in the capillary fringe above the alluvial ground-water table. These phreatophytic cottonwoods would be highly dependent upon stream flow and vulnerable to declining river flows due to river regulation or climate change.     

The dependence of fish community structure and dynamics on floodplain and riparian ecotone zone in Parana River,Brazil

Flood intensity was a main factor determining access to the riparian/floodplain ecotonal resources of the upper Parana River, Brazil, and in consequence determining yield of the dominant trophic groups, which are fish feeding on flooded grasslands and on riparian fruits and leaves. Prey and predator density both declined in years of low floods, and predators did not recover until the next year of high flood, more slowly than in the case prey fish, most probably due to differences in life cycle length. The life cycle of one of the most important species, Curimba (Prochilodus lineatus — Characidae) depends on access to the floodplain lagoons and return to the reservoir after maturation for intensive growth. The riparian/floodplain habitat complexity and connectivity has great significance for fish community dynamics and fisheries yield, and may serve a reason to create a national park in the last floodplain section of the upper Parana River.     

Mechanisms of Riparian Cottonwood Decline Along Regulated Rivers

Decline of riparian forests has been attributed to hydrologic modifications to river flows. However, little is known about the physiological and structural adjustments of riparian forests subject to modified flow regimes, and the potential for forest restoration using historic flow regimes is poorly understood. In this paired river study, we compared hydrology, water relations, and forest structure in cottonwood-dominated floodplains of the regulated Green River to those of the unregulated Yampa River. We measured floodplain groundwater levels, soil water availability, cottonwood xylem pressure (Ψ_xp), and leaf-level stomatal conductance (g_s) to assess current impacts of river regulation on the water status of adult cottonwoods. We also simulated a flood on the former floodplain of the regulated river to evaluate its impact on cottonwood water relations. Canopy and root structure were quantified with estimates of cottonwood leaf area and percent live canopy and root density and biomass, respectively. Regulation of the Green River has lowered the annual peak flow yet raised minimum flows in most years, resulting in a 60% smaller stage change, and lowered soil water availability by as much as 70% compared to predam conditions. Despite differences in water availability, daily and seasonal trends in Ψ_xp and g_s were similar for cottonwoods on the regulated and unregulated rivers. In addition, soil water added with the experimental flood had little effect on cottonwood water relations, contrary to our expectations of alleviated water stress. Green River cottonwoods had 10%–30% lower stand leaf area, 40% lower root density, and 25% lower root biomass compared with those for Yampa River cottonwoods. Our results suggest that water relations at the leaf and stem level are currently similar for Yampa and Green River trees due to structural adjustments of cottonwood forests along the Green River, triggered by river regulation.     

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.     

High‐resolution riparian vegetation mapping to prioritize conservation and restoration in an impaired desert river

In highly impaired watersheds, it is critical to identify both areas with desirable habitat as conservation zones and impaired areas with the highest likelihood of improvement as restoration zones. We present how detailed riparian vegetation mapping can be used to prioritize conservation and restoration sites within a riparian and instream habitat restoration program targeting 3 native fish species on the San Rafael River, a desert river in southeastern Utah, United States. We classified vegetation using a combination of object‐based image analysis (OBIA) on high‐resolution (0.5 m), multispectral, satellite imagery with oblique aerial photography and field‐based data collection. The OBIA approach is objective, repeatable, and applicable to large areas. The overall accuracy of the classification was 80% (Cohen's κ = 0.77). We used this high‐resolution vegetation classification alongside existing data on habitat condition and aquatic species' distributions to identify reaches' conservation value and restoration potential to guide management actions. Specifically, cottonwood (Populus fremontii) and tamarisk (Tamarix ramosissima) density layers helped to establish broad restoration and conservation reach classes. The high‐resolution vegetation mapping precisely identified individual cottonwood trees and tamarisk thickets, which were used to determine specific locations for restoration activities such as beaver dam analogue structures in cottonwood restoration areas, or strategic tamarisk removal in high‐density tamarisk sites. The site prioritization method presented here is effective for planning large‐scale river restoration and is transferable to other desert river systems elsewhere in the world.     

Forest Development, Wood Jams, and Restoration of Floodplain Rivers in the Puget Lowland, Washington

Historically in Puget Lowland rivers, wood jams were integral to maintaining an anastomosing channel pattern and a dynamic channel–floodplain connection; they also created deep pools. In the late 1800s, wood was removed from most rivers, rivers were isolated from their floodplains, and riparian forests were cut down, limiting wood recruitment. An exception to this history is an 11-km-long reach of the Nisqually River, which has natural banks and channel pattern and a mature floodplain forest. We use field and archival data from the Nisqually River to explore questions relevant to restoring large rivers in the Pacific Northwest and other forested temperate regions. In particular, we focus on the relation between recovery of in-channel wood accumulations and valley bottom forest conditions and explore implications for river restoration strategies. We find that restoring large rivers depends on establishing riparian forests that can provide wood large enough to function as key pieces in jams. Although the frequency of large trees in the Nisqually valley bottom in 2000 is comparable with that of 1873 land surveys, many formerly more abundant Thuja plicata (western red cedar) were cut down in the late 1800s, and now hardwoods, including Populus trichocarpa (black cottonwood) and Acer macrophyllum (bigleaf maple), are also abundant. Pseudotsuga menziesii (Douglas fir) and fast-growing P. trichocarpa commonly form key pieces that stabilize jams, suggesting that reforested floodplains can develop naturally recruited wood jams within 50 to 100 years, faster than generally assumed. Based on the dynamic between riparian forests, wood recruitment, and wood jams in the Nisqually River, we propose a planning framework for restoring self-sustaining dynamic river morphology and habitat to forested floodplain rivers.     

Nitrogen cycling in two riparian forest soils under different geomorphic conditions

At the floodplain scale, spatial pattern and successional development of riparian vegetation are under the control of geomorphic processes. The geomorphic and hydraulic characteristics of stream channels affect the sorting of organic material and inorganic sediment through erosion/sedimentation during floods. In turn, the proportion of fine sediments fractions differs by location within a given community of riparian forest succession. In this paper we tested the effect of geomorphic features of floodplains, through soil grain size sorting, on the nitrogen cycling in riparian forest soils. Two typical riparian forests formed by vertical accretion deposits from repeated addition of sediments from overbank flow have been chosen along the River Garonne, southwest France. These riparian forests had equivalent vegetation, flood frequency and duration, differing only in soil grain size composition: one riparian forest had sandy soils and the other had loamy soils. The evolution of the main soil physical and chemical parameters as well as denitrification (DNT), N uptake (N _U ) and mineralization (N _M ) rates were measured monthly over a period of 13 months in the two study sites. The loamy riparian forest presented a better physical retention of suspended matter during floods. Moreover,in situ denitrification rates (DNT) and N uptake by plants (N _U ) measured in the loamy riparian forest soils were significantly greater than in the sandy soils. Although DNT and N _U could be in competition for available nitrogen, the peak rates of these two processes did not occur at the same period of the year, N _U being more important during the dry season when DNT was minimum, while DNT rates were maximum following the spring floods. N retention by uptake (N _U ) and loss by DNT represented together the equivalent of 32% of total organic nitrogen deposited during floods on the sandy riparian forest soils and 70% on the loamy ones. These significant differences between the two sites show that, at the landscape level, one should not estimate the rates of N _U and DNT, in riparian forests soils only on the basis of vegetation, but should take also into account the geomorphic features of the floodplain.     

Can environmental flows moderate riparian invasions? The influence of seedling morphology and density on scour losses in experimental floods

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Relative contributions of sexual and asexual regeneration strategies in Populus nigra and Salix alba during the first years of establishment on a braided gravel bed river

Populus nigra L. var betulifolia and Salix alba L. var alba are early successional riparian tree species threatened throughout Continental Europe by significant changes to the natural physical processes governing their natural habitat – geomorphologically active floodplains. River management activities have dramatically altered natural patterns of river flow and rates of sediment delivery along rivers, with possible consequences for the balance between sexual and asexual regeneration strategies in these species. Conservation strategies will benefit from a greater understanding of the ways in which dynamic physical processes on the floodplain influence sexual and asexual recruitment. This paper describes a field survey investigating the relative abundance and spatial distributions of P. nigra and S. alba sexual and asexual recruits during the first years of establishment along a braided gravel bed river. Sexual and asexual recruits were identified by excavation along transects in a wet and a dry field season and distributional differences were described in terms of elevation on the floodplain, local sediment type and exposure to floodwaters. Regeneration was overwhelmingly from seed in the first 2–3 years following recruitment, but poor survival rates among sexual recruits saw a shift in the relative abundance of regeneration strategies over time. In relating hydrological data to recruitment, unseasonal flood disturbances had a negative effect on recruitment from seed and a positive effect on vegetative regeneration. Seedlings were associated with fine sediment deposits and were restricted primarily to low elevations on the flood plain, while asexual recruits had a wider spatial distribution. Certain microsite types were unique to either regeneration strategy.     

Determining factors of cottonwood planting survival in a desert river restoration project

Planting native riparian trees can help recover wildlife and fish habitat on a local scale, when full recovery of natural processes that sustain riparian ecosystems is infeasible. To help improve planting success, we determined which environmental factors and management practices most influenced survival of planted Populus fremontii (Fremont cottonwood) in a field experiment on the San Rafael River, Utah, U.S.A. We planted 474 approximately 2‐m‐tall trees and tracked survival for 1.25 years. We used logistic regression to evaluate whether tree height, elevation above the river channel, distance to existing cottonwood or Salix exigua (coyote willow), soil conductivity, soil texture, planting depth, planting method (mechanical auger vs. hand‐digging), and provision of natural and commercial supplements affected survival probability. Survival probability decreased with elevation above the river channel bottom and was greater in auger‐dug than hand‐dug holes. Survival probability was lower in soils with the highest salinity levels and was lower in sandy soils than soils with silt and clay. Survival may be improved by planting well above the channel to avoid flooding impacts but within 2 m above the channel in auger‐dug holes to ensure access to soil moisture. Testing soil salinity and texture in areas with suitable elevation could also help improve survival. Approximately 35% of trees survived to the end of the study period, indicating that planting can help recover riparian habitat locally, especially if survival is improved in future planting efforts. However, full recovery of desired riparian habitat throughout the floodplain will require natural flows.     

Evapotranspiration at the land/water interface in a semi-arid drainage basin

1. Evapotranspiration (ET) is a major source of water depletion from riverine systems in arid and semiarid climates. Water budgets have produced estimates of total depletions from riparian vegetation ET for a 320‐km reach of the Middle Rio Grande, New Mexico, U.S.A., that have ranged from 20 to 50% of total depletions from the river. 2. Tower‐based micrometeorological measurements of riparian zone ET throughout the growing season using three‐dimensional eddy covariance provided high quality estimates of ET at the stand scale. 3. A dense stand of salt cedar (111–122 cm year^–1) and a mature cottonwood (Populus deltoides ssp. wislizenia Eckenwelder) stand with an extensive understory of salt cedar (Tamaria ramosissima Ledeb) and Russian olive (Eleagnus angustifolia L.) (123 cm year^–1) had the highest rates of annual ET. A mature cottonwood stand with a closed canopy had intermediate rates of ET (98 cm year^–1). A less dense salt cedar stand had the lowest rates of ET (74–76 cm year^–1). 4. Summer leaf area index (LAI) measurements within the four stands were positively correlated with daily ET rates. LAI measurements throughout the growing season coupled to riparian vegetation classification is a promising method for improving riverine corridor estimates of total annual riparian zone ET along a reach of river. 5. Combining recent estimates of the extent of riparian vegetation along the 320 km length of the Middle Rio Grande, from Landsat 7 imagery with annual growing season measurements of ET at the four riparian stands yields a first‐order riverine corridor estimate of total riparian zone ET of 150–250 × 10⁶ m³ year^–1. This is approximately 20–33% of total estimated depletions along this reach of river.     

Differing influences of natural and artificial disturbances on riparian cottonwoods from prairie to mountain ecoregions in Alberta, Canada

Aim Ecoregions represent biophysical zones where environmental factors enable the development of particular plant communities. Ecoregions are generally large but abrupt transitions occur in areas with rapid physical change. A particularly abrupt transitional sequence occurs in the Rocky Mountain region of south‐western Alberta where fescue prairie, aspen parkland and mountain ecoregions occur within 15 km. To investigate plant adaptation across ecoregions, our study investigated the influences of a natural disturbance (flooding) and an artificial disturbance (cattle grazing) on reproductive and population processes of black cottonwood (Populus balsamifera subsp. trichocarpa, Torr. & Gray), the dominant riparian tree. Location We studied cottonwoods throughout their elevational range along two free‐flowing, first‐order streams, Yarrow and Drywood creeks. Cottonwood was the only prominent tree in the prairie ecoregion, the dominant riparian tree in the parkland and extended upward through the montane ecoregion where it was a pioneer species for the mixed coniferous–deciduous woodland. Cottonwoods did not occur in the higher elevation sub‐alpine ecoregion. Methods Thirty‐six cross‐sectional sampling transects were located across the three ecoregions with cottonwoods, and in ungrazed and grazed areas of each ecoregion. Rectangular 100 m² tree and 2 m² seedling quadrats were positioned along the transects, and substrate and vegetation were assessed. Historic hydrological data were analysed relative to flood recurrences and seasonal flow patterns. Results Overall, the cottonwoods displayed a sawtooth shaped ‘punctuated progressive age structure’ with many young trees, progressively fewer older trees, and about four pulses of increased recruitment over the past century. This was considered to provide a healthy cottonwood population and recruitment pulses were apparently associated with flood events with appropriate peak timing and magnitude and a gradual post‐flood stage recession. However, analyses of tree, sapling and seedling data indicated that flood‐associated seedling recruitment was less important and clonal processes were more important for cottonwood recruitment in the montane ecoregion, the highest ecoregion with cottonwoods. The correlation between flood events and cottonwood recruitment was strongest in the mid‐elevation parkland ecoregion suggesting greater reliance on flood‐associated seedling recruitment. There was little correlation with flooding and limited recruitment in the fescue prairie ecoregion in recent decades and the disturbed age structure probably results from cattle impacts that have prevented recruitment and produced a decrepit cottonwood forest population. Main conclusions These analyses suggested that a healthy cottonwood population displayed a sawtooth shaped ‘punctuated progressive age structure’ and that cottonwood reproduction processes varied across ecoregions with increased clonality in the highest montane ecoregion. Cattle grazing impacts on reproduction were most severe in the lowest prairie ecoregion that is treeless except for the riparian zone. We conclude that appropriate strategies of instream flow regulation, land‐use policies and practices, and conservation and restoration efforts should be refined according to ecoregion to recognize the differences in cottonwood reproductive and population ecology.     

Avian Density and Nest Survival on the San Pedro River: Importance of Vegetation Type and Hydrologic Regime

Abstract: Lowland riparian vegetation in the southwestern United States is critically important for maintaining a high richness and density of breeding birds. Further investigation is needed within riparian corridors, however, to evaluate the relative importance of vegetation type and hydrologic regime for avian density and nest survival as targets for regional conservation or restoration efforts. We estimated the densities of 40 bird species and for species grouped on the basis of nest height and dependence on surface water in gallery cottonwood–willow (Populus spp.–Salix spp.) forests, saltcedar (Tamarix spp.) shrub lands, and terrace vegetation types along a gradient in the hydrologic regime of the San Pedro River, Arizona, USA. We also assessed nest survival for shrub-nesting insectivores and herbivores. Canopy-nesting birds as a group and 14 individual bird species reached their greatest densities in cottonwood forests regardless of the hydrologic regime. Water-dependent birds as a group reached their highest density in both intermittent- and perennial-flow cottonwood stands, but certain species occurred almost exclusively in perennial-flow sites. Two shrub-nesting species and the brown-headed cowbird (Molothrus ater) were most abundant in saltcedar shrub lands, and the brown-headed cowbird was most abundant in saltcedar stands with intermittent flows. Mesquite (Prosopis spp.) and big sacaton (Sporobolus wrightii) grassland each maintained the highest densities of certain species within ≥1 hydrologic regime. Shrub-nesting insectivores had the greatest nest survival in cottonwood, including Arizona Bell's vireo (Vireo bellii arizonae), and also had lower proportions of nests parasitized and preyed upon, although 95% confidence intervals among vegetation types overlapped. Nest survival for both shrub-nesting insectivores and herbivores was lowest in intermittent-flow saltcedar, although, again, confidence intervals overlapped. Nest survival was lower in parasitized than nonparasitized nests in mesquite and across vegetation types for Arizona Bell's vireo and in cottonwood for Abert's towhee (Pipilo aberti). Riparian management that maintains heterogeneous riparian vegetation types, including floodplain vegetation comprising cottonwood–willow gallery riparian forests with some stretches of perennial flow, are important for maintaining the high diversity and abundance of breeding birds on the San Pedro River and probably across the region. Cottonwood stands also appear to maintain highest nest survival for some shrub-nesting birds.     

Effects of flooding on leaf development,transpiration, and photosynthesis in narrowleaf cottonwood,a willow-like poplar

The narrowleaf cottonwood, Populus angustifolia, occurs in occasionally flooded, low elevation zones along river valleys near the North American Rocky Mountains. This small poplar has narrow leaves and fine branching and thus resembles willows, which are commonly flood-tolerant. We investigated the flood response of narrowleaf cottonwoods and a related native hybrid, jackii cottonwood (P. × jackii = P. balsamifera × P. deltoides), by studying saplings of 24 clones in a greenhouse, with some pots being inundated to provide the flood treatment. Flooding slightly reduced leaf numbers (−10%), and leaf sizes were reduced by about 21% in female P. angustifolia versus a 50% reduction in the female hybrids. Flooding-reduced stomatal conductance and net photosynthetic rate, and reduced transpiration particularly in P. × jackii. The effects on foliar gas exchange declined over a 5-week interval, suggesting compensation. The moderate impact of flooding supports the hypothesis that narrowleaf cottonwoods are flood-tolerant, and we anticipate that these trees could provide traits to increase the flood tolerance of fast-growing hybrid poplars. The results further indicate that female cottonwoods may be more flood-tolerant than males, and females could be more successful in lower, flood-prone sites.     

Recruitment limitation along disturbance gradients in river flood plains

Abstract. Question: Along river floodplains lower distribution limits of plant species seem largely determined by their tolerance to rarely occurring floods in the growing season. Such distribution patterns remain fixed for many years suggesting additional effects of winter floods at lower positions. Our objective was to investigate the direct and indirect effects of winter floods on colonization of floodplains in a series of field experiments. Location: River Rhine, The Netherlands. Methods: We measured the direct effects of winter floods on seedling survival and seed removal and survival at low and high floodplain elevation. Indirect effects of winter flooding through changes in the soil were investigated by measuring seedling emergence on soil transplants that were exchanged between high and low floodplain elevation. To investigate indirect effects of floods on the germination environment through changes in the vegetation structure, we measured the effects of vegetation removal on recruitment of sown species. Results: Recruitment was seed limited at both floodplain elevations. An additional effect of vegetation removal on seedling emergence was also observed. Soil types from both zones did not differently affect seedling emergence. Seeds were not removed from the soil surface by a single winter flood. Moreover, seeds remained viable in the soil for at least two years, while the experimental plots were flooded several times during the experimental period. During one of those floods a thick sand layer was deposited at the low zone and subsequently no seedlings were observed anymore. Conclusions: Colonization of low floodplain zones in years between subsequent summer floods is prevented by seed limitation while the direct effects of winter floods are limited except for irregularly occurring sand depositions.     

Growth of riparian cottonwoods: a developmental pattern and the influence of geomorphic context

Cottonwoods are poplars (Populus sp.) adapted to riparian (streamside) zones and an understanding of their growth within these zones will assist with river management for cottonwood conservation and in the recognition of superior parental genotypes for hybrid poplar breeding programs. In this study we analyzed cottonwood growth in native riparian zones and compared growth along three study reaches of the Oldman River in Alberta, Canada that differed in geomorphic context, particularly the extent that the river channel was constrained by steep banks and bedrock. We used dendrochronology to analyze trunk growth patterns, and measured annual radial increments (RI) and basal area increments (BAI) of 278 narrowleaf cottonwoods (P. angustifolia), black cottonwoods (P. trichocarpa), their intrasectional hybrids, and natural intersectional hybrids with prairie cottonwoods (P. deltoides). The trees displayed common growth patterns with four phases: (I) a 3–7-year establishment phase with RI of about 1–2 mm/year, (II) a growth acceleration phase of about 15 years with RI increasing to the (III) RI growth peak of about 3 mm/year, and then (IV) the mature growth phase with relatively constant BAI and progressively declining RI. This general pattern was consistent across study reaches but the durations and growth rates of the phases differed along with forest stand structure. Along the unconstrained alluvial reach with a broad floodplain and dynamic channel, extensive and dense forest groves occurred. This increased tree competition, as evidenced by reduced RI and BAI in the mature phase. In contrast, along a constrained reach trees were restricted to sparse, narrow bands and their increased growth rates in the mature phase probably reflected reduced competition. Cottonwoods along the intermediate reach demonstrated an intermediate combination of forest and growth characteristics. Genotypic effects were slight although P. angustifolia had reduced RI during the establishment phase. These results demonstrate that within native riparian zones cottonwoods display an inherent growth pattern that reflects the trees' life history, and that growth rates and transitions are influenced by the geomorphic context that influences forest structure.     

Resizing a River: A Downscaled,Seasonal Flow Regime Promotes Riparian Restoration

While riverine organisms are adapted to the natural flow regime, it is impractical to fully restore natural flows along most regulated rivers. We propose an alternative with the delivery of downscaled flow regimes that provide the seasonal patterns that are essential for aquatic and riparian ecosystems. The Bridge River in British Columbia provided a novel case study as a downscaled flow regime commenced in 2000 along a reach that had generally experienced no flow for the prior half‐century. The experimental flow delivered a mean discharge of about 3 m³/s, versus the pre‐dam mean of 100 m³/s, with a seasonal pattern that mimicked the natural snowmelt‐dominated pattern. To assess the environmental response, we investigated black cottonwoods, Populus trichocarpa, the dominant riparian trees, in the pre‐flow versus post‐flow intervals, using tree ring interpretation for growth analyses and age determination. Sparse mature trees established prior to the 1948 damming did not show significant growth changes in the pre‐ versus post‐flow intervals. In contrast, younger trees that established closer to the river in the decade prior to 2000 displayed significant growth increases by 2002, and juveniles established after 2000 demonstrated faster initial growth than juveniles established before 2000. Further, bands of cottonwood saplings resulted from seedling recruitment along the new river fringe, particularly in 2002, 2003, and 2004, years with gradual flow recession. These responses demonstrate that a downscaled, seasonal flow regime provided environmental benefit, thereby restoring some river function and resulting in a resized river flanked by narrow and reproducing cottonwood bands.