A multi-scale assessment of the occurrence of exotic plants on the Olympic Peninsula,Washington

Abstract. Exotic plants were surveyed in 208 plots within the Dungeness and Hoh river watersheds on the Olympic Peninsula, Washington, USA. Landscape patch types included uplands (clearcuts, young and mature forests) and riparian zones (cobble bars, shrub patches, riparian forests, and alder flats). Patterns of exotic plant invasions were assessed between watersheds, between riparian and upland areas, among patch types, and within clearcuts. 52 exotic plant species were encountered, accounting for 23% of the flora in each watershed. In both watersheds, exotic species richness was approximately 33% greater in riparian zones than in uplands, and mean number and cover of exotic species were > 50% greater in riparian zones than in uplands. Among landscape patch types, richness and mean number and cover of exotics was highest in young riparian patches, intermediate in clearcuts and riparian forests, and lowest in young and mature forests. The exception to this was Hoh alder flats, which had the highest mean cover of exotic plants. Cover of exotic plants peaked in uplands 3 to 7 yr after clearcutting, then decreased with increased canopy closure. Disturbance type and time since disturbance were major factors influencing invasibility. Landscape patch size, position within watershed (distance from patch to human population centers, major highway, or river mouth), and environmental variables (slope, aspect, and elevation) were not important indicators of landscape patch invasibility. Riparian zones facilitated movement of exotic plants through landscapes, but did not appear to act as sources of exotic plants for undisturbed upland areas.     

Patterns of Plant Invasions: A Case Example in Native Species Hotspots and Rare Habitats

Land managers require landscape-scale information on where exotic plant species have successfully established, to better guide research, control, and restoration efforts. We evaluated the vulnerability of various habitats to invasion by exotic plant species in a 100,000 ha area in the southeast corner of Grand Staircase-Escalante National Monument, Utah. For the 97 0.1-ha plots in 11 vegetation types, exotic species richness (log₁₀) was strongly negatively correlated to the cover of cryptobiotic soil crusts (r = −0.47, P < 0.001), and positively correlated to native species richness (r = 0.22, P < 0.03), native species cover (r = 0.23, P < 0.05), and total nitrogen in the soil (r = 0.40, P < 0.001). Exotic species cover was strongly positively correlated to exotic species richness (r = 0.68, P < 0.001). Only 6 of 97 plots did not contain at least one exotic species. Exotic species richness was particularly high in locally rare, mesic vegetation types and nitrogen rich soils. Dry, upland plots (n = 51) had less than half of the exotic species richness and cover compared to plots (n = 45) in washes and lowland depressions that collect water intermittently. Plots dominated by trees had significantly greater native and exotic species richness compared to plots dominated by shrubs. For the 97 plots combined, 33% of the variance in exotic species richness could be explained by a positive relationship with total plant cover, and negative relationships with the cover of cryptobiotic crusts and bare ground. There are several reasons for concern: (1) Exotic plant species are invading hot spots of native plant diversity and rare/unique habitats. (2) The foliar cover of exotic species was greatest in habitats that had been invaded by several exotic species.(3) Continued disturbance of fragile cryptobiotic crusts by livestock, people, and vehicles may facilitate the further invasion of exotic plant species. This revised version was published online in July 2006 with corrections to the Cover Date.     

How Does Restoration of Native Canopy Affect Understory Vegetation Composition? Evidence from Riparian Communities of the Hunter Valley Australia

Restoration of native vegetation often focuses on the canopy layer species, with the assumption that regeneration of the understory elements will occur as a consequence. The goal of this study was to assess the influence of canopy restoration on the composition and abundance of understory plant species assemblages along riparian margins in the Hunter Valley, NSW, Australia. We compared the floristic composition (richness, abundance, and diversity) of understory species between nonrevegetated (open) and canopy revegetated plots across five sites. A number of other factors that may also influence understory vegetation, including soil nutrients, proximity to main channel, and light availability, were also measured. We found that sites where the canopy had been restored had lower exotic species richness and abundance, as well as higher native species cover, but not native species richness, compared with open sites. Multivariate analysis of plots based on plant community composition showed that revegetated sites were associated with lower total species diversity, light availability, and exotic cover. This study has found that the restoration of the canopy layer does result in lower exotic species richness and cover, and higher native species cover and diversity in the understory, a desirable restoration outcome. Our results provide evidence that restoration of native canopy species may facilitate restoration of native understory species; however, other interventions to increase native species richness of the understory should also be considered as part of management practice.     

Linear declines in exotic and native plant species richness along an increasing altitudinal gradient in the Snowy Mountains,Australia

Abstract Declines in plant species richness with increasing altitude are common, but the form of the relationship can vary, with both monotonic decreasing relationships and humped relationship recorded. However, these different richness to altitude relationships may be due to methods that used different plot sizes/areas and survey efforts. To explore native and exotic plant richness along an altitudinal gradient in the Snowy Mountains of Australia, we consistently surveyed plots that were 120 m² in area at 39 sites ranging from 540 to 2020 m. To relate exotic plant richness to disturbance, we surveyed plots at 16 sites along main roads and 23 sites along minor roads and also compared these 39 roadside plots to 120‐m² plots located in undisturbed vegetation adjacent to the roadside (native plant richness was only surveyed in 25 of these 39 adjacent plots). We found a negative linear relationship between total, exotic and native species richness and altitude for plots on the side of main roads (16 sites) and minor roads (23 sites). For adjacent plots negative linear relationships were significant for all measures of species richness except for native species adjacent to major roads. As the pattern occurred for exotics it is less likely to be due to historical constraints on the species pools. The pattern could be influenced by difference in levels of disturbance along the gradient, although any such gradient in disturbance would have to apply to roadside and adjacent plots on major and minor roads. Therefore, it may be due to other factors such as changes in climate along the altitudinal gradient, although additional sampling including direct measures of climatic conditions, soil and disturbance factors would be needed to determine if this was the case.     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999     

Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico

This article presents an analysis of plant species richness and diversity and its association with climatic and soil variables along a 1300‐m elevation gradient on the Cerro Tláloc Mountain in the northern Sierra Nevada in Mexico. Two 1000‐m² tree sampling plots were created at each of 21 selected sampling sites, as well as two 250‐m² plots for shrubs and six 9‐m² plots for herbaceous plants. Species richness and diversity were estimated for each plant life form, and beta diversity between sites was estimated along the gradient. The relationship between species richness and diversity and environmental variables was modelled using simple linear correlation and regression trees. Species richness and diversity showed a unimodal pattern with a bias towards high values in the lower half of the elevation gradient under study. This response was consistent for all three life forms. Beta diversity increased steadily along the elevation gradient, being lower between contiguous sites at intermediate elevations and high – the species replacement rate was nearly 100%– between sites at the extremes of the gradient. Few species were adapted to the full spectrum of environmental variation along the elevation gradient studied. The regression tree suggests that differences in species richness are mainly influenced by elevation (temperature and humidity) and soil variables, namely A₂ permanent wilting point, organic matter and horizon field capacity and A₁ horizon Mg²⁺.     

Long-term responses in population dynamics and diversity of small mammals in riparian and upland habitats within an agricultural landscape

Riparian zones in agricultural landscapes provide linear non-crop habitats for a variety of plant and mammal species, and hence are an important component of biodiversity. To date, variable responses of abundance, species richness, and species diversity of small mammals have been recorded in riparian and upland habitats. To address this variability, we provide a detailed analysis of seasonal changes in abundance and diversity of terrestrial small-mammal communities over a 7-year period within an agricultural landscape in south-central British Columbia, Canada. We tested the hypotheses (H) that abundance, species richness, and species diversity of communities of small mammals (H₁), and demographic parameters of reproduction, recruitment, and survival of the major species: deer mouse (Peromyscus maniculatus) and montane vole (Microtus montanus) (H₂), would be higher in riparian than upland habitats. Mean total abundance of small mammals was higher in summer and winter, and species richness higher in summer, in riparian than hedgerow habitats. Winter population data supported the total and species abundance patterns for small mammals, but species richness was similar, and diversity lower, in riparian than hedgerow sites during winter periods. Deer mice were the dominant species in terms of abundance and reproductive output for pregnancies and recruitment, but not survival, in riparian sites. Montane voles were similar in abundance and demographic parameters in the two habitats. House mice (Mus musculus) preferred hedgerows and wandering shrews (Sorex vagrans) riparian sites. Demographic parameters for deer mice and montane voles indicated that both riparian and hedgerow sites were “source” rather than “sink” habitats, and likely contribute to maintenance of mammal diversity in agricultural landscapes.     

Relationship between native and exotic plant species at multiple savannah sites

We tested whether the recently proposed two‐part measure of degree of invasion (DI) of a community relating exotic proportion of cover to exotic proportion of richness can characterize patterns of plant invasions at multiple savannah sites in Southern Africa. Regression analysis was performed on transformed data to assess how this two‐part measure of DI compares to other metrics of community invasibility. The results indicate that at the plot level, the absolute cover of exotics was not significantly related to native cover for three sites out of four assessed (R² ≤ 0.17; P > 0.05). Also, at all four sites, no significant relationships were detected between native and exotic plant richness at both the 1‐m² and 400‐m² plot scales. By contrast, significant (P < 0.05) positive linear relationships were observed between exotic proportion of richness and exotic proportion of cover at all sites (R² was as high as 0.67 and 0.97 for two sites). Our results indicate that the new two‐part measure of DI is able to characterize patterns of plant invasions across plant communities in African savannahs.     

Long‐term effects of exclusion of grazing stock on degraded herbaceous plant communities in a riparian Eucalyptus camaldulensis forest in south‐eastern Australia

Abstract Stock grazing has degraded many riparian ecosystems around the world. However, the potential for ecosystem recovery following the removal of grazing stock is poorly known. We developed a conceptual model to predict the responses of native and exotic herbaceous plants to grazing exclusion, based on site productivity and the degree of initial vegetation degradation. The effects of excluding grazing stock on richness, cover and composition of herbaceous plants were examined over 12 years in the degraded understorey of a riparian forest in Gulpa Island State Forest in south‐eastern Australia. We predicted that grazing exclusion would lead to limited changes in vegetation cover, richness and composition, owing to presumed low site productivity and the high degree of understorey degradation. Results showed that the cover, richness and composition of native and exotic species varied significantly among years. Over all plots, regions and years, total cover was slightly but significantly lower in grazed than in ungrazed plots (43.4% vs. 50.8%). While the cover of native plants increased over time in both treatments, the rate of increase was slightly greater in ungrazed plots. Grazing exclusion had no effect on the richness of native or exotic species, but had a significant but minor impact on plant composition, with different common species (mostly exotics) being promoted or diminished in ungrazed plots. The composition of grazed and ungrazed areas did not become more different over time. Overall, the effects that could be attributed to grazing exclusion were relatively minor and transient. Results are consistent with predictions based on site productivity and initial degradation, and should not be extrapolated to other more productive, or less degraded, riparian systems.     

Rio Grande Wild Turkey Habitat Selection in the Southern Great Plains

ABSTRACT We recorded telemetry locations from 1,129 radiotagged turkeys (Meleagris gallopavo intermedia) on 4 study areas in the Texas Panhandle and southwestern Kansas, USA, from 2000 to 2004. Analyses of telemetry locations indicated both sexes selected riparian vegetative zones. Females did not select grazed or nongrazed pastures for daily movements. However, females did select nongrazed pastures for nest sites on 2 study areas and males selected for grazed pastures at one study area during the breeding season. We compared nest sites (n = 351) to random sites using logistic regression, which indicated height of visual obstruction, percent canopy cover, and percent bare ground provided the highest predictive power (P ≤ 0.003) for characteristics describing nest-site selection. Nest-site vegetative characteristics between vegetative zones differed primarily in composition: upland zone nest sites had more (P ≤ 0.001) shrubs and riparian zone nest sites had more (P ≤ 0.001) grass. There were no differences in measured nest site vegetative characteristics between pasture types, but there were differences between available nesting cover in grazed and nongrazed pastures. Random plots in grazed pastures had less grass cover (P ≤ 0.001) and more bare ground (P = 0.002). Because of cattle impacts on average grass height and availability, grazing would likely have the highest impact on nesting in riparian zones due to turkey use of grass as nesting cover. An appropriate grazing plan to promote Rio Grande turkey nesting habitat would include grazing upland zones in the spring, when it likely has little impact on nesting-site selection, and grazing riparian zones following breeding season completion. Grazing at light to moderate intensities with periods of rest did not affect male turkey pasture use and may have continued to maintain open areas used by male turkeys for displaying purposes.     

Effects of proximity to riparian zones on avian species richness and abundance in montane aspen woodlands

Riparian zones often provide more food or nesting resources than surrounding ecosystems and thus support more species or a greater abundance of birds. However, the extent to which the positive effects of riparian zones extend into adjoining habitats has rarely been investigated. We examined bird species richness and abundance in aspen (Populus tremuloides) woodlands of the upper East River Valley in Colorado, a region with extensive stands of aspen located upslope from riparian zones dominated by willows (Salix spp.). Our objective was to determine how species richness and abundance in aspen stands located closer to riparian zones compared to that of aspen stands farther away. To control for elevation effects, we conducted point counts for birds and examined the effects of riparian proximity on avian species richness and abundance in three elevation belts (low, 2840–2875 m; middle, 2940–2960 m; and high, 3040–3060 m). We determined riparian proximity by measuring distances from each census site to the nearest body of water and to the nearest patch of willow. Proximity to major willow patches (≥6500 m²) had a stronger influence on species richness and abundance than did proximity to water or smaller patches of willows. Total species richness and abundance significantly increased with proximity to major willow patches at low elevation sites only. This relationship was driven by a greater prevalence of riparian‐nesting species at lower elevation sites, where aspens were generally closer to riparian zones. The positive effects of willows diminished with abrupt changes in elevation. Our results highlight the importance of protecting willows around riparian zones in valley bottoms where habitat destruction caused by human land use is most prominent.     

Plant biomass and species composition along an environmental gradient in montane riparian meadows

In riparian meadows, narrow zonation of the dominant vegetation frequently occurs along the elevational gradient from the stream edge to the floodplain terrace. We measured plant species composition and above- and belowground biomass in three riparian plant communities—a priori defined as wet, moist, and dry meadow—along short streamside topographic gradients in two montane meadows in northeast Oregon. The objectives were to: (1) compare above- and belowground biomass in the three meadow communities; (2) examine relations among plant species richness, biomass distribution, water table depth, and soil redox potential along the streamside elevational gradients. We installed wells and platinum electrodes along transects (perpendicular to the stream; n=5 per site) through the three plant communities, and monitored water table depth and soil redox potential (10 and 25 cm depth) from July 1997 to August 1999. Mean water table depth and soil redox potential differed significantly along the transects, and characterized a strong environmental gradient. Community differences in plant species composition were reflected in biomass distribution. Highest total biomass (live+dead) occurred in the sedge-dominated wet meadows (4,311±289 g/m²), intermediate biomass (2,236±221 g/m²) was seen in the moist meadow communities, dominated by grasses and sedges, and lowest biomass (1,403±113 g/m²) was observed in the more diverse dry meadows, dominated by grasses and forbs. In the wet and moist communities, belowground biomass (live+dead) comprised 68–81% of the totals. Rhizome-to-root ratios and distinctive vertical profiles of belowground biomass reflected characteristics of the dominant graminoid species within each community. Total biomass was positively correlated with mean water table depth, and negatively correlated with mean redox potential (10 cm and 25 cm depths; P <0.01) and species richness (P <0.05), indicating that the distribution of biomass coincided with the streamside edaphic gradient in these riparian meadows.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Life-history Habitat Matching in Invading Non-native Plant Species

We briefly reviewed the literature on habitat matching in invading non-native plant species. Then we hypothesized that the richness and cover of native annual and perennial plant species integrate complex local information of vegetation and soils that would help to predict invasion success by similarly adapted non-native plant species. We tested these ‘life-history habitat matching’ relationships in 603 0.1-ha plots, including 294 plots in Colorado, which were relatively high for the cover of native perennial plant species, and for 309 0.1-ha plots in southern Utah, which were relatively high in the cover of native annual plant species. We found strong positive relationships between the richness and foliar cover for both native and non-native species, whether they were annual or perennial species (0.34 > r ² < 0.53; P < 0.0001). We also found significant positive relationships between the cover of native annual species at a site and the richness (r ² = 0.13; P < 0.0001) and the foliar cover (r ² = 0.06; P < 0.0001) of non-native annual species. The proportion of non-native annual species in the flora of a plot also increased significantly with the foliar cover of native annual species. Conversely, the richness and cover of non-native annual species were significantly negatively associated with the foliar cover of native perennial species (r ² = 0.05 and 0.06, respectively; P < 0.0001). The cover of non-native annual or perennial species was not significantly correlated with soil texture variables, %N, or %C. We conclude that there may be a high degree of life-history habitat matching by non-native annual species in these study sites. Information on native annual and perennial species richness and cover may help characterize the complex soils, climate, and disturbance environment in which similarly adapted non-native plant species establish and gain foliar cover.     

Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA

It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40–50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r ²=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r ² _adjacent=0.72, r ² _distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.     

Effects of elevated nitrogen and exotic plant invasion on soil seed bank composition in Joshua Tree National Park

We investigated the effects of exotic species invasion and 3?years of nitrogen (N) fertilization on the soil seed bank in Joshua Tree National Park, California, USA at four sites along an N deposition gradient. We compared seed bank composition and density in control (no N added) and fertilized (30?kg?N?ha^?1?year^?1) plots to determine if the seed bank would reflect aboveground changes due to N fertilization. Soil samples were collected and germinated in a greenhouse over 2?years. In the field, invasive species cover responded positively to N fertilization. However, we did not observe increased seed density of exotic invasive species in fertilized plots. While no significant differences were detected between treatments within sites, exotic invasive grass seeds overwhelmed the seed bank at all sites. Significant differences between sites were found, which may be due to differences in level of invasion, historic N deposition, and soil surface roughness. Sites experiencing low N deposition had the highest seed bank species richness for both control and fertilized treatments. Aboveground plant density did not correlate well with seed bank density, possibly due to the inherent patchiness of soil seed banks and differential ability of species to form seed banks. This seed bank study provided insight into site-specific impacts on native versus invasive species composition of soil seed banks, as well as magnitude of invasion and restoration potential at invaded sites.     

Encroachment of upland Mediterranean plant species in riparian ecosystems of southern Portugal

Riparian ecosystems have unique biodiversity, are highly sensitive to disturbance and anthropogenic influence. As world water resources become scarcer, scientists predict greater competition among species for water resources. Indeed, increased encroachment of upland plants into the riparian zone is already occurring, irreversibly changing riparian plant communities. Since semi-arid regions such as Mediterranean-type ecosystems are likely to follow this same trajectory, assessing the contributions of riparian versus upland (sclerophyllous) plants to community composition is important. A survey of seventy 2 km-long riparian transects on the Sado and Guadiana watersheds in southern Portugal assessed (1) the woody riparian plant community composition, (2) how much richness is due to strictly riparian plants versus sclerophyllous upland plants, and (3) which combinations of biotic and abiotic factors allow higher species richness in the strictly riparian, sclerophyllous, and overall plant communities. The survey detected 53 different woody plant species (28 endemic) across all communities. Riparian community richness was on average 16 species, seven of which were strictly riparian and the remainder being sclerophyllous, exotic species or fruit trees. Sclerophyllous plant species occurred consistently across sampling units (90% of transects). On average, 46% of the total woody plant community richness was due to strictly riparian plants and 28% was due to sclerophyllous plants. Community richness was positively affected by the area of shrubs in the riparian zone and by the absence of human activities and goats. Surrounding landscape pattern only affected the strictly riparian plant richness. These results suggest that natural and human-mediated disturbances in riparian ecosystems create gaps and clearings for which riparian and sclerophyllous plants compete. Establishment success seems to be related to the propagule pressure of the neighbouring landscape, its diversity and density, as well as the presence of herbivores. Preserving strictly riparian plants, removing exotic species, preventing grazing, and promoting riparian values (recreation, aesthetics and the provision of ecosystem services) will aid the future conservation of the unique biodiversity of riparian ecosystems.     

Effects of past logging and grazing on understory plant communities in a montane Colorado forest

Throughout Pinus ponderosa–Pseudotsuga menziesii forests of the southern Colorado Front Range, USA, intense logging and domestic grazing began at the time of Euro-American settlement in the late 1800s and continued until the early 1900s. We investigated the long-term impacts of these settlement-era activities on understory plant communities by comparing understory composition at a historically logged and grazed site to that of an environmentally similar site which was protected from past use. We found that species richness and cover within functional groups rarely differed between sites in either upland or riparian areas. Multivariate analyses revealed little difference in species composition between sites on uplands, though compositional differences were apparent in riparian zones. Our findings suggest that settlement-era logging and grazing have had only minor long-term impacts on understories of upland Front Range P. ponderosa–P. menziesii forests, though they have had a greater long-term influence on riparian understories, where these activities were likely the most intense. This article was written and prepared by US Government employees on official time, and therefore it is in the public domain and not subject to copyright.     

The Goldilocks effect: intermittent streams sustain more plant species than those with perennial or ephemeral flow

1. Drylands worldwide are typified by extreme variability in hydrologic processes, which structures riparian communities at various temporal and spatial scales. One key question is how underlying differences in hydrology over the length of interrupted perennial rivers influence spatial and temporal patterns in species richness and species composition. 2. We examined effects of differences in dry season hydrology on species richness, composition and cover of herbaceous plant communities in the streamside zone (the zone influenced directly by low flows in the channel). Data were collected at ephemeral, intermittent and perennial flow reaches on three rivers of the desert Southwest (Arizona, U.S.A.): Lower Cienega Creek, Hassayampa River and Lower San Pedro River. 3. Patterns of species richness varied with temporal scale of analysis, that is between single‐year and multi‐year time frames. At the annual timescale, quadrat species richness (m^?2) and herbaceous cover were higher at sites with perennial flow than at either intermittent or ephemeral sites. In contrast to this single‐year pattern, the highest long‐term richness occurred at intermittent sites. 4. Quadrat species richness, total species richness at a site (per 18 1‐m² plots) and cover were more variable year to year at non‐perennial sites than at perennial flow sites. On two of the three rivers, ephemeral sites had the highest inter‐annual compositional variance, while the perennial sites had the lowest. 5. Compositional differences between the hydrologic site types were dominated by species turnover, not nestedness. The perennial sites had more wetland and perennial species than the other two site types. The intermittent sites had more annual species than did the other two types. 6. High long‐term species richness and distinct species composition of intermittent sites are probably sustained by pronounced temporal variability in environmental conditions (i.e. frequent and persistent flow events, and dry periods). Plants at these sites take advantage of greater moisture than those at ephemeral sites and also experience less competition from resident species than those at perennial sites. 7. Conservation of desert riparian diversity depends upon the protection of consistently wet conditions at perennial flow sites, as well as the maintenance of the processes that cause fluctuations in environmental conditions at non‐perennial sites.     

A Modified-Whittaker nested vegetation sampling method

A standardized sampling technique for measuring plant diversity is needed to assist in resource inventories and for monitoring long-term trends in vascular plant species richness. The widely used Whittaker plot (Shmida 1984) collects species richness data at multiple spatial scales, using 1 m², 10 m², and 100 m² subplots within a 20 m × 50 m (1000 m²) plot, but it has three distinct design flaws involving the shape and placement of subplots. We modified and tested a comparable sampling design (Modified-Whittaker plot) that minimizes the problems encountered in the original Whittaker design, while maintaining many of its attractive attributes. We overlaid the two sampling methods in forest and prairie vegetation types in Larimer County, Colorado, USA (n=13 sites) and Wind Cave National Park, South Dakota, USA (n=19 sites) and showed that the modified design often returned significantly higher (p<0.05) species richness values in the 1 m², 10 m², and 100 m² subplots. For all plots, except seven ecotone plots, there was a significant difference (p<0.001) between the Whittaker plot and the Modified-Whittaker plot when estimating the total number of species in the 1000 m² plots based on linear regressions of the subplot data: the Whittaker plot method, on average, underestimated plant species richness by 34%. Species-area relationships, using the Modified-Whittaker design, conformed better to published semilog relationships, explaining, on average, 92% of the variation. Using the original Whittaker design, the semilog species-area relationships were not as strong, explaining only 83% of the variation, on average. The Modified-Whittaker plot design may allow for better estimates of mean species cover, analysis of plant diversity patterns at multiple spatial scales, and trend analysis from monitoring a series of strategically-placed, long-term plots.     

Impacts of mixed severity wildfire on exotic plants in a Colorado ponderosa pine–Douglas-fir forest

The 2002 Hayman Fire burned with mixed severity across 55,800 ha of montane Colorado forest, including pre-existing plots that were originally measured for understory plant composition and cover in 1997. We examined the influence of the Hayman Fire on exotic plants by remeasuring these plots annually from 2003 to 2007. We found that (1) exotic richness and cover generally increased as fire severity and time since fire increased; (2) the exotic species present in a plot before the fire were also largely present in the plot postfire, regardless of fire severity; (3) most of the new postfire species in a plot were present elsewhere in the study area before the fire, although some new species were truly new invaders that were not found in prefire surveys; (4) lightly burned riparian forests were not more susceptible to exotic invasion than surrounding uplands that burned with similar severity; and (5) native and exotic richness and cover were positively correlated or uncorrelated for all fire severities and years. Our findings indicate that exotics were stimulated by the Hayman Fire, especially in severely burned areas. However, exotic richness and cover remain low as of 2007, and correlations between native and exotic richness and cover suggest that exotics have not yet interfered with native understory development. Therefore, we conclude that exotic plants are not a major ecological threat at present, but recommend that monitoring be continued to evaluate if they will pose a threat in future years.