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.     

Can bryophytes be used to characterize hydrologic permanence in forested headwater streams?

Recent court cases have questioned whether all headwater streams, particularly those that are not perennial, fall within the protective boundaries of the Clean Water Act. Rapid field-based indicators of hydrologic permanence are critically needed for jurisdictional determinations. The study objective was to determine whether characteristics of bryophyte assemblages in forested headwater streams can be useful indicators of hydrologic permanence. We sampled bryophytes from the channel thalweg at 113 sites across 10 forests in the U.S. which varied in hydrologic permanence. Hydrologic permanence was based on hydrologic status during spring (wet season) and late summer (dry season) visits. Perennial sites had flow (surface or visibly interstitial) during both spring and summer visits. Intermittent sites flowed during spring but were either dry or had surface water limited to isolated pools in summer. Ephemeral sites did not have flow during either visit. The species composition significantly differed between ephemeral and perennial stream reaches. The species composition of intermittent streams overlapped with both ephemeral and perennial streams. Three species and six families were identified as indicators of particular permanence classes. Liverworts occurred more frequently at perennial than at intermittent or ephemeral sites. Bryophytes with cushion and turf growth forms were common to ephemeral sites, whereas mat and weft forms were more common to perennial sites. Acrocarpous mosses were more frequent at ephemeral than at perennial sites. The frequencies of bryophyte-moisture associations (derived from the literature) across the three permanence categories were significantly different. Species associated with dry habitats were more frequently encountered at ephemeral than perennial sites, whereas species associated with wet habitats were more frequent at perennial than ephemeral sites. Species richness varied among forests and permanence classes. Ephemeral sites tended to have higher species richness than perennial sites; however, this pattern did not hold for all forests. Bryophytes should be included among the tools for jurisdictional determinations, much like vascular plants are used to help delineate wetlands. Growth forms and higher level taxonomy of bryophytes (i.e., phyla) can be particularly useful for making jurisdictional determinations because of their ability to discriminate ephemeral from intermittent and perennial reaches and they are measures that can be rapidly trained and routinely used by agencies making such determinations.     

Drying of a temperate,intermittent river has little effect on adjacent riparian arthropod communities

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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.     

Return of Fire to a Free-Flowing Desert River: Effects on Vegetation

After a long period in which fuel loads were sparse, fire recently has occurred with high frequency in the ungrazed riparian zone of the Upper San Pedro River in southern Arizona's Chihuahuan Desert. We studied four accidental fires that occurred during 1994–2003 (two in different years at the same site). Woody vegetation was contrasted between three burned sites and matched spatial controls, and before and after the most recent fire. Herbaceous vegetation was sampled in multiple years producing a chronosequence of time since fire (from 4 months to 8 years). Riparian fire was associated with reductions in woody plant species diversity and canopy cover. In contrast, fire caused a short-term (2 year) pulse of herbaceous plant diversity, driven by annual species, and persistent increase in herbaceous cover. Path analysis indicated that the increase in herbaceous cover was mediated in part by the reduction in tree canopy cover. Ordination (nonmetric multidimensional scaling) and regression analysis also indicated that canopy cover and/or fire played a role in structuring the herbaceous community, although its effects were secondary to that of hydrologic factors (stream flow rate, seasonal flood size). By converting riparian forests to grasslands and savannahs, fire may be shifting structure of the Upper San Pedro floodplain vegetation closer toward conditions present during past centuries when fire was frequent in the upland desert grasslands and embedded riparian corridor.     

The underappreciated role of life history in mediating the functional consequences of biodiversity change

Biodiversity is changing on both global and local scales, motivating research to understand the consequences of these changes for how communities and ecosystems function. Here, we explore the role of life history strategies in mediating biodiversity and ecosystem functioning. In particular, we evaluate how the composition, biomass (% cover), and richness of perennial (persistence ≥ 1 year) and ephemeral (persistence < 1 year) species change along a gradient of increasing seaweed species richness on a rocky shoreline. We show that the majority of biomass is comprised of perennial species, especially where overall richness is low, whereas the majority of species are ephemeral, especially where overall richness is high. We then present and discuss the results of an 18‐month field manipulation quantifying the factorial effects of tidal elevation, wave exposure, herbivore removals, thermal and desiccation stress amelioration, and nutrient additions on perennial versus ephemeral species. In particular, the diversity of ephemeral species was strongly affected, relative to perennial species, by tidal elevation, wave exposure, and herbivory; herbivores reduced diversity of ephemeral species relative to perennials. Relative to perennial cover, ephemeral cover was greater higher on the shore, in more wave‐exposed habitats, and where herbivores were removed, plots were unscreened, and/or nutrients were added. Thus, perennials and ephemerals responded differently to environmental context and experimental manipulation. We compared nitrate uptake and photosynthesis rates of ephemeral and perennial species and found that maximum nitrate uptake and photosynthesis rates of ephemerals were twice as high as those of perennials. These results highlight the disproportionate roles that ephemeral species play in mediating ecosystem‐level processes. In combination with our comparisons of the diversity and cover of perennial and ephemeral species along a biodiversity gradient, these results demonstrate the utility of incorporating life history traits into our efforts to understand the functional consequences of biodiversity change.     

Does stream flow structure woody riparian vegetation in subtropical catchments?

The primary objective of this study was to test the relevance of hydrological classification and class differences to the characteristics of woody riparian vegetation in a subtropical landscape in Queensland, Australia. We followed classification procedures of the environmental flow framework ELOHA – Ecological Limits of Hydrologic Alteration. Riparian surveys at 44 sites distributed across five flow classes recorded 191 woody riparian species and 15, 500 individuals. There were differences among flow classes for riparian species richness, total abundance, and abundance of regenerating native trees and shrubs. There were also significant class differences in the occurrence of three common tree species, and 21 indicator species (mostly native taxa) further distinguished the vegetation characteristics of each flow class. We investigated the influence of key drivers of riparian vegetation structure (climate, depth to water table, stream‐specific power, substrate type, degree of hydrologic alteration, and land use) on riparian vegetation. Patterns were explained largely by climate, particularly annual rainfall and temperature. Strong covarying drivers (hydrology and climate) prevented us from isolating the independent influences of these drivers on riparian assemblage structure. The prevalence of species considered typically rheophytic in some flow classes implies a more substantial role for flow in these classes but needs further testing. No relationships were found between land use and riparian vegetation composition and structure. This study demonstrates the relevance of flow classification to the structure of riparian vegetation in a subtropical landscape, and the influence of covarying drivers on riparian patterns. Management of environmental flows to influence riparian vegetation assemblages would likely have most potential in sites dominated by rheophytic species where hydrological influences override other controls. In contrast, where vegetation assemblages are dominated by a diverse array of typical rainforest species, and other factors including broad‐scale climatic gradients and topographic variables have greater influence than hydrology, riparian vegetation is likely to be less responsive to environmental flow management.     

Do intermittent and ephemeral Mediterranean rivers belong to the same river type?

The benthic macroinvertebrate communities and ecological quality of eleven temporary rivers (seven intermittent and four ephemeral) in Dadia National Park, north-eastern Greece, were examined with respect to the degree of flow temporality. Sampling took place during the high flow season at both ephemeral and intermittent sites and during the low flow season only at the intermittent ones, which receded to pools. Despite the remarkable seasonal variation in both the hydrology and ecology of the intermittent rivers, the various metrics and indices as well as the multivariate analyses confirmed the clear distinction between the two river types (ephemeral and intermittent). Existing European quality indices do not sufficiently differentiate between ephemeral and intermittent river types, and thus cannot reliably discriminate the degree of natural variability from human induced stressors in temporary rivers.     

Arbuscular mycorrhizal fungi associated with Populus-Salix stands in a semiarid riparian ecosystem

This study examined the activity, species richness, and species composition of the arbuscular mycorrhizal fungal (AMF) community of Populus-Salix stands on the Verde River (Arizona, USA), quantified patterns of AMF richness and colonization along complex floodplain gradients, and identified environmental variables responsible for structuring the AMF community. Samples from 61 Populus-Salix stands were analyzed for AMF and herbaceous composition, AMF colonization, gravimetric soil moisture, soil texture, per cent organic matter, pH, and concentrations of nitrate, bicarbonate phosphorus and exchangeable potassium. AMF species richness declined with stand age and distance from and elevation above the channel and was positively related to perennial species cover and richness and gravimetric soil moisture. Distance from and elevation above the active channel, forest age, annual species cover, perennial species richness, and exchangeable potassium concentration all played a role in structuring the AMF community in this riparian area. Most AMF species were found across a wide range of soil conditions, but a subset of species tended to occur more often in hydric areas. This group of riparian affiliate AMF species includes several not previously encountered in the surrounding Sonoran desert.     

Spatial and temporal patterns of species richness in a riparian landscape

Aim To test for control of vascular plant species richness in the riparian corridor by exploring three contrasting (although not mutually exclusive) hypotheses: (1) longitudinal patterns in riparian plant species richness are governed by local, river‐related processes independent of the regional species richness, (2) riparian plant species richness is controlled by dispersal along the river (longitudinal control), and (3) the variation in riparian plant species richness mirrors variation in regional richness (lateral control). Location The riparian zones of the free‐flowing Vindel River and its surrounding river valley, northern Sweden. Methods We used data from three surveys, undertaken at 10‐year intervals, of riparian reaches (200‐m stretches of riverbank) spanning the entire river. In addition, we surveyed species richness of vascular plants in the uplands adjacent to the river in 3.75‐km² large plots along the same regional gradient. We explored the relationship between riparian and upland flora, and various environmental variables. We also evaluated temporal variation in downstream patterns of the riparian flora. Results Our results suggest that local species richness in boreal rivers is mainly a result of local, river‐related processes and dispersal along the corridor. The strongest correlation between species richness and the environment was a negative one between species number and soil pH, but pH varied within a narrow range. We did not find evidence for a correlation between species richness on regional and local scales. We found that the local patterns of species richness for naturally occurring vascular plants were temporally variable, probably in response to large‐scale disturbance caused by extreme floods. Most previous studies have found a unimodal pattern of species richness with peaks in the middle reaches of a river. In contrast, on two of three occasions corresponding to major flooding events, we found that the distribution of species richness of naturally occurring vascular plants resembled that of regional diversity: a monotonic decrease from headwater to coast. We also found high floristic similarity between the riparian corridor and the surrounding landscape. Main conclusions These results suggest that local processes control patterns of riparian species richness, but that species composition is also highly dependent on the regional species pool. We argue that inter‐annual variation in flood disturbance is probably the most important factor producing temporal variability of longitudinal species richness patterns.