Barbara E. Ralston

Riparian vegetation, Colorado River, and climate: Five decades of spatiotemporal dynamics in the Grand Canyon with river regulation

Documentation of the interacting effects of river regulation and climate on riparian vegetation has typically been limited to small segments of rivers or focused on individual plant species. We examine spatiotemporal variability in riparian vegetation for the Colorado River in Grand Canyon relative to river regulation and climate, over the five decades since completion of the upstream Glen Canyon Dam in 1963. Long-term changes along this highly modified, large segment of the river provide insights for management of similar riparian ecosystems around the world. We analyze vegetation extent based on maps and imagery from eight dates between 1965 and 2009, coupled with the instantaneous hydrograph for the entire period. Analysis confirms a net increase in vegetated area since completion of the dam. Magnitude and timing of such vegetation changes are river stage-dependent. Vegetation expansion is coincident with inundation frequency changes and is unlikely to occur for time periods when inundation frequency exceeds approximately 5%. Vegetation expansion at lower zones of the riparian area is greater during the periods with lower peak and higher base flows, while vegetation at higher zones couples with precipitation patterns and decreases during drought. Short pulses of high flow, such as the controlled floods of the Colorado River in 1996, 2004, and 2008, do not keep vegetation from expanding onto bare sand habitat. Management intended to promote resilience of riparian vegetation must contend with communities that are sensitive to the interacting effects of altered flood regimes and water availability from river and precipitation.

Do beavers promote the invasion of non-native Tamarix in the Grand Canyon riparian zone?

Beavers (Castor canadensis Kuhl) can influence the competitive dynamics of plant species through selective foraging, collection of materials for dam creation, and alteration of hydrologic conditions. In the Grand Canyon National Park, the native Salix gooddingii C.R.Ball (Goodding’s willow) and Salix exigua Nutt. (coyote willow) are a staple food of beavers. Because Salix competes with the invasive Tamarix ramosissima Ledeb., land mangers are concerned that beavers may cause an increase in Tamarix through selective foraging of Salix. A spatial analysis was conducted to assess whether the presence of beavers correlates with the relative abundance of Salix and Tamarix. These methods were designed to detect a system-wide effect of selective beaver foraging in this large study area (367 linear km of riparian habitat). Beavers, Salix, and Tamarix co-occurred at the broadest scales because they occupied similar riparian habitat, particularly geomorphic reaches of low and moderate resistivity. Once the affinity of Salix for particular reach types was accounted for, the presence of Salix was independent of beaver distribution. However, there was a weak positive association between beaver presence and Salix cover. Salix was limited to geomorphic settings with greater sinuosity and distinct terraces, while Tamarix occurred in sinuous and straighter sections of river channel (cliffs, channel margins) where it dominated the woody species composition. After accounting for covariates representing river geomorphology, the proportion of riparian surfaces covered by Tamarix was significantly greater for sites where beavers were present. This indicates that either Tamarix and beavers co-occur in similar habitats, beavers prefer habitats that have high Tamarix cover, or beavers contribute to Tamarix dominance through selective use of its native woody competitors. The hypothesis that beaver herbivory contributes to Tamarix dominance should be considered further through more mechanistic studies of beaver foraging processes and longterm plant community response.

Variation in species-level plant functional traits over wetland indicator status categories

Abstract Wetland indicator status (WIS) describes the habitat affinity of plant species and is used in wetland delineations and resource inventories. Understanding how species‐level functional traits vary across WIS categories may improve designations, elucidate mechanisms of adaptation, and explain habitat optima and niche. We investigated differences in species‐level traits of riparian flora across WIS categories, extending their application to indicate hydrologic habitat. We measured or compiled data on specific leaf area (SLA), stem specific gravity (SSG), seed mass, and mature height of 110 plant species that occur along the Colorado River in Grand Canyon, Arizona. Additionally, we measured leaf δ13C, δ15N, % carbon, % nitrogen, and C/N ratio of 56 species with C3 photosynthesis. We asked the following: (i) How do species‐level traits vary over WIS categories? (ii) Does the pattern differ between herbaceous and woody species? (iii) How well do multivariate traits define WIS categories? (iv) Which traits are correlated? The largest trait differences among WIS categories for herbaceous species occurred for SSG, seed mass, % leaf carbon and height, and for woody species occurred for height, SSG, and δ13C. SSG increased and height decreased with habitat aridity for both woody and herbaceous species. The δ13C and hence water use efficiency of woody species increased with habitat aridity. Water use efficiency of herbaceous species increased with habitat aridity via greater occurrence of C4 grasses. Multivariate trait assemblages differed among WIS categories. Over all species, SLA was correlated with height, δ13C, % leaf N, and C/N; height was correlated with SSG and % leaf C; SSG was correlated with % leaf C. Adaptations of both herbaceous and woody riparian species to wet, frequently inundated habitats include low‐density stem tissue. Adaptations to drier habitats in the riparian zone include short, high‐density cavitation‐resistant stem tissue, and high water use efficiency. The results enhance understanding about using traits to describe plant habitat in riparian systems.

Changes in Community-Level Riparian Plant Traits over Inundation Gradients, Colorado River, Grand Canyon

Comparisons of community-level functional traits across environmental gradients have potential for identifying links among plant characteristics, adaptations to stress and disturbance, and community assembly. We investigated community-level variation in specific leaf area (SLA), plant mature height, seed mass, stem specific gravity (SSG), relative cover of C4 species, and total plant cover over hydrologic zones and gradients in years 2013 and 2014 in the riparian plant community along the Colorado River in the Grand Canyon. Vegetation cover was lowest in the frequently inundated active channel zone, indicating constraints on plant establishment and production by flood disturbance and anaerobic stress. Changes in trait values over hydrologic zones and inundation gradients indicate that frequently inundated plots exhibit a community-level ruderal strategy with adaptation to submergence (high SLA and low SSG, height, seed mass, C4 relative cover), whereas less frequently inundated plots exhibit adaptation to drought and infrequent flood disturbance (low SLA and high SSG, height, seed mass, C4 relative cover). Variation in traits not associated with inundation suggests niche differentiation and multiple modes of community assembly. The results enhance understanding of future responses of riparian communities of the Grand Canyon to anticipated drying and changes in hydrologic regime.

Functional traits and ecological affinities of riparian plants along the Colorado River in Grand Canyon

Abstract. Trait-based approaches to vegetation analyses are becoming more prevalent in studies of riparian vegetation dynamics, including responses to flow regulation, groundwater pumping, and climate change. These analyses require species trait data compiled from the literature and floras or original field measurements. Gathering such data makes trait-based research time intensive at best and impracticable in some cases. To support trait-based analysis of vegetation along the Colorado River through Grand Canyon, a data set of 20 biological traits and ecological affinities for 179 species occurring in that study area was compiled. This diverse flora shares species with many riparian areas in the western USA and includes species that occur across a wide moisture gradient. Data were compiled from published scientific papers, unpublished reports, plant fact sheets, existing trait databases, regional floras, and plant guides. Data for ordinal environmental tolerances were more readily available than were quantitative traits. More publicly available data are needed for traits of both common and rare southwestern U.S. plant species to facilitate comprehensive, traitbased research. The trait data set is free to use and can be downloaded from ScienceBase: https://www.sciencebase.gov/catalog/item/58af41dee4b01ccd54f9f2ff and https://dx.doi.org/10.5066/F7QV3JN1

Taxonomic and Compositional Differences of Ground-Dwelling Arthropods in Riparian Habitats in Glen Canyon, Arizona, USA

Abstract. The disturbance history, plant species composition, productivity, and structural complexity of a site can exert bottom-up controls on arthropod diversity, abundance, and trophic structure. Regulation alters the hydrology and disturbance regimes of rivers and affects riparian habitats by changing plant quality parameters. Fifty years of regulation along the Colorado River downstream of Glen Canyon Dam has created a no-analog, postdam “lower” riparian zone close to the waters edge that includes tamarisk (Tamarix sp.), a nonnative riparian shrub. At the same time, the predam “upper” facultative riparian zone has persisted several meters above the current flood stage. In summer 2009, we used pitfall traps within these 2 riparian zones that differ in plant composition, productivity, and disturbance frequency to test for differences in arthropod community (Hymenoptera, Arachnida, and Coleoptera) structure. Arthropod community structure differed substantially between the 2 zones. Arthropod abundance and species richness was highest in the predam upper riparian zone, even though there was a greater amount of standing plant biomass in the postdam lower riparian zone. Omnivore abundance was proportionately greater in the upper riparian zone and was associated with lower estimated productivity values. Predators and detritivores were proportionately greater in the postdam lower riparian zone. In this case, river regulation may create habitats that support species of spiders and carabid beetles, but few other species that are exclusive to this zone. The combined richness found in both zones suggests a small increase in total richness and functional diversity for the Glen Canyon reach of the Colorado River.