Thomas J. Stohlgren

EXOTIC PLANT SPECIES INVADE HOT SPOTS OF NATIVE PLANT DIVERSITY

Some theories and experimental studies suggest that areas of low plant spe- cies richness may be invaded more easily than areas of high plant species richness. We gathered nested-scale vegetation data on plant species richness, foliar cover, and frequency from 200 1-m 2 subplots (20 1000-m 2 modified-Whittaker plots) in the Colorado Rockies (USA), and 160 1-m 2 subplots (16 1000-m 2 plots) in the Central Grasslands in Colorado, Wyoming, South Dakota, and Minnesota (USA) to test the generality of this paradigm. At the 1-m 2 scale, the paradigm was supported in four prairie types in the Central Grasslands, where exotic species richness declined with increasing plant species richness and cover. At the 1-m 2 scale, five forest and meadow vegetation types in the Colorado Rockies contradicted the paradigm; exotic species richness increased with native-plant species richness and foliar cover. At the 1000-m 2 plot scale (among vegetation types), 83% of the variance in exotic species richness in the Central Grasslands was explained by the total percentage of nitrogen in the soil and the cover of native plant species. In the Colorado Rockies, 69% of the variance in exotic species richness in 1000-m 2 plots was explained by the number of native plant species and the total percentage of soil carbon. At landscape and biome scales, exotic species primarily invaded areas of high species richness in the four Central Grasslands sites and in the five Colorado Rockies vegetation types. For the nine vegetation types in both biomes, exotic species cover was positively correlated with mean foliar cover, mean soil percentage N, and the total number of exotic species. These patterns of invasibility depend on spatial scale, biome and vegetation type, spatial autocorrelation effects, availability of resources, and species-specific responses to grazing and other disturbances. We conclude that: (1) sites high in herbaceous foliar cover and soil fertility, and hot spots of plant diversity (and biodiversity), are invasible in many landscapes; and (2) this pattern may be more closely related to the degree resources are available in native plant communities, independent of species richness. Exotic plant in- vasions in rare habitats and distinctive plant communities pose a significant challenge to land managers and conservation biologists.

THE INVASION PARADOX: RECONCILING PATTERN AND PROCESS IN SPECIES INVASIONS

The invasion paradox describes the co-occurrence of independent lines of support for both a negative and a positive relationship between native biodiversity and the invasions of exotic species. The paradox leaves the implications of native-exotic species richness relationships open to debate: Are rich native communities more or less susceptible to invasion by exotic species? We reviewed the considerable observational, experimental, and theoretical evidence describing the paradox and sought generalizations concerning where and why the paradox occurs, its implications for community ecology and assembly processes, and its relevance for restoration, management, and policy associated with species invasions. The crux of the paradox concerns positive associations between native and exotic species richness at broad spatial scales, and negative associations at fine scales, especially in experiments in which diversity was directly manipulated. We identified eight processes that can generate either negative or positive native-exotic richness relationships, but none can generate both. As all eight processes have been shown to be important in some systems, a simple general theory of the paradox, and thus of the relationship between diversity and invasibility, is probably unrealistic. Nonetheless, we outline several key issues that help resolve the paradox, discuss the difficult juxtaposition of experimental and observational data (which often ask subtly different questions), and identify important themes for additional study. We conclude that natively rich ecosystems are likely to be hotspots for exotic species, but that reduction of local species richness can further accelerate the invasion of these and other vulnerable habitats.

The rich get richer: patterns of plant invasions in the United States

Observations from islands, small-scale experiments, and mathematical models have generally supported the paradigm that habitats of low plant diversity are more vulnerable to plant invasions than areas of high plant diversity. We summarize two independent data sets to show exactly the opposite pattern at multiple spatial scales. More significant, and alarming, is that hotspots of native plant diversity have been far more heavily invaded than areas of low plant diversity in most parts of the United States when considered at larger spatial scales. Our findings suggest that we cannot expect such hotspots to repel invasions, and that the threat of invasion is significant and predictably greatest in these areas.

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.

HOW GRAZING AND SOIL QUALITY AFFECT NATIVE AND EXOTIC PLANT DIVERSITY IN ROCKY MOUNTAIN GRASSLANDS

We used multiscale plots to sample vascular plant diversity and soil characteristics in and adjacent to 26 long-term grazing exclosure sites in Colorado, Wyoming, Montana, and South Dakota, USA. The exclosures were 7–60 yr old (31.2 ± 2.5 yr, mean ± 1 se). Plots were also randomly placed in the broader landscape in open rangeland in the same vegetation type at each site to assess spatial variation in grazed landscapes. Consistent sampling in the nine National Parks, Wildlife Refuges, and other management units yielded data from 78 1000-m2 plots and 780 1-m2 subplots. We hypothesized that native species richness would be lower in the exclosures than in grazed sites, due to competitive exclusion in the absence of grazing. We also hypothesized that grazed sites would have higher native and exotic species richness compared to ungrazed areas, due to disturbance (i.e., the intermediate-disturbance hypothesis) and the conventional wisdom that grazing may accelerate weed invasion. Both hypotheses were soundly rej...

Riparian zones as havens for exotic plant species in the central grasslands

In the Central Grasslands of the United States, we hypothesized that riparian zones high in soil fertility would contain more exotic plant species than upland areas of low soil fertility. Our alternate hypothesis was that riparian zones high in native plant species richness and cover would monopolize available resources and resist invasion by exotic species. We gathered nested-scale vegetation data from 40 1 m2subplots (nested in four 1000 m2 plots) in both riparian and upland sites at four study areas in Colorado, Wyoming, and South Dakota (a total of 320 1 m2 subplots and 32 1000 m2 plots). At the 1 m2 scale, mean foliar cover of native species was significantly greater (P<0.001) in riparian zones (36.3% ± 1.7%) compared to upland sites (28.7% ± 1.5%), but at this small scale there were no consistent patterns of native and exotic species richness among the four management areas. Mean exotic species cover was slightly higher in upland sites compared to riparian sites (9.0% ± 3.8% versus 8.2% ± 3.0% cover). However, mean exotic species richness and cover were greater in the riparian zones than upland sites in three of four management areas. At the 1000 m2 scale, mean exotic species richness was also significantly greater (P<0.05) in riparian zones (7.8 ± 1.0 species) compared to upland sites (4.8 ± 1.0 species) despite the heavy invasion of one upland site. For all 32 plots combined, 21% of the variance in exotic species richness was explained by positive relationships with soil % silt (t =1.7, P=0.09) and total foliar cover (t = 2.4, P=0.02). Likewise, 26% of the variance in exotic species cover (log10 cover) was explained by positive relationships with soil % silt (t =2.3, P=0.03) and total plant species richness (t = 2.5, P=0.02). At landscape scales (four 1000 m2 plots per type combined), total foliar cover was significantly and positively correlated with exotic species richness (r=0.73, P<0.05) and cover (r=0.74, P<0.05). Exotic species cover (log10 cover) was positively correlated with log10% N in the soil (r=0.61, P=0.11) at landscape scales. On average, we found that 85% (±5%) of the total number of exotic species in the sampling plots of a given management area could be found in riparian zones, while only 50% (±8%) were found in upland plots. We conclude that: (1) species-rich and productive riparian zones are particularly invasible in grassland ecosystems; and (2) riparian zones may act as havens, corridors, and sources of exotic plant invasions for upland sites and pose a significant challenge to land managers and conservation biologists.

Evidence that local land use practices influence regional climate, vegetation, and stream flow patterns in adjacent natural areas

We present evidence that land use practices in the plains of Colorado influence regional climate and vegetation in adjacent natural areas in the Rocky Mountains in predictable ways. Mesoscale climate model simulations using the Colorado State University Regional Atmospheric Modelling System (RAMS) projected that modifications to natural vegetation in the plains, primarily due to agriculture and urbanization, could produce lower summer temperatures in the mountains. We corroborate the RAMS simulations with three independent sets of data: (i) climate records from 16 weather stations, which showed significant trends of decreasing July temperatures in recent decades; (ii) the distribution of seedlings of five dominant conifer species in Rocky Mountain National Park, Colorado, which suggested that cooler, wetter conditions occurred over roughly the same time period; and (iii) increased stream flow, normalized for changes in precipitation, during the summer months in four river basins, which also indicates cooler summer temperatures and lower transpiration at landscape scales. Combined, the mesoscale atmospheric/land-surface model, short-term trends in regional temperatures, forest distribution changes, and hydrology data indicate that the effects of land use practices on regional climate may overshadow larger-scale temperature changes commonly associated with observed increases in CO2 and other greenhouse gases.

GLOBAL POTENTIAL NET PRIMARY PRODUCTION PREDICTED FROM VEGETATION CLASS, PRECIPITATION, AND TEMPERATURE

Net primary production (NPP), the difference between CO2 fixed by photosynthesis and CO2 lost to autotrophic respiration, is one of the most important components of the carbon cycle. Our goal was to develop a simple regression model to estimate global NPP using climate and land cover data. Approximately 5600 global data points with observed mean annual NPP, land cover class, precipitation, and temperature were compiled. Precipitation was better correlated with NPP than temperature, and it explained much more of the variability in mean annual NPP for grass- or shrub-dominated systems (r2 = 0.68) than for tree-dominated systems (r2 = 0.39). For a given precipitation level, tree-dominated systems had significantly higher NPP (approximately 100-150 g C m(-2) yr(-1)) than non-tree-dominated systems. Consequently, previous empirical models developed to predict NPP based on precipitation and temperature (e.g., the Miami model) tended to overestimate NPP for non-tree-dominated systems. Our new model developed at the National Center for Ecological Analysis and Synthesis (the NCEAS model) predicts NPP for tree-dominated systems based on precipitation and temperature; but for non-tree-dominated systems NPP is solely a function of precipitation because including a temperature function increased model error for these systems. Lower NPP in non-tree-dominated systems is likely related to decreased water and nutrient use efficiency and higher nutrient loss rates from more frequent fire disturbances. Late 20th century aboveground and total NPP for global potential native vegetation using the NCEAS model are estimated to be approximately 28 Pg and approximately 46 Pg C/yr, respectively. The NCEAS model estimated an approximately 13% increase in global total NPP for potential vegetation from 1901 to 2000 based on changing precipitation and temperature patterns.

SPATIAL HETEROGENEITY INFLUENCES NATIVE AND NONNATIVE PLANT SPECIES RICHNESS

Spatial heterogeneity may have differential effects on the distribution of native and nonnative plant species richness. We examined the effects of spatial heterogeneity on native and nonnative plant species richness distributions in the central part of Rocky Mountain National Park, Colorado, USA. Spatial heterogeneity around vegetation plots was characterized using landscape metrics, environmental/topographic variables (slope, aspect, elevation, and distance from stream or river), and soil variables (nitrogen, clay, and sand). The landscape metrics represented five components of landscape heterogeneity and were measured at four spatial extents (within varying radii of 120, 240, 480, and 960 m) using the FRAGSTATS landscape pattern analysis program. Akaikes Information Criterion adjusted for small sample size (AICc) was used to select the best models from a set of multiple linear regression models developed for native and nonnative plant species richness at four spatial extents and three levels of ecological hierarchy (i.e., landscape, land cover, and community). Both native and nonnative plant species richness were positively correlated with edge density, Simpsons diversity index and interspersion/juxtaposition index, and were negatively correlated with mean patch size. The amount of variation explained at four spatial extents and three hierarchical levels ranged from 30% to 70%. At the landscape level, the best models explained 43% of the variation in native plant species richness and 70% of the variation in nonnative plant species richness (240-m extent). In general, the amount of variation explained was always higher for nonnative plant species richness, and the inclusion of landscape metrics always significantly improved the models. The best models explained 66% of the variation in nonnative plant species richness for both the conifer land cover type and lodgepole pine community. The relative influence of the components of spatial heterogeneity differed for native and nonnative plant species richness and varied with the spatial extent of analysis and levels of ecological hierarchy. The study offers an approach to quantify spatial heterogeneity to improve models of plant biodiversity. The results demonstrate that ecologists must recognize the importance of spatial heterogeneity in managing native and nonnative plant species.

Potential habitat distribution for the freshwater diatom Didymosphenia geminata in the continental US

The diatom Didymosphenia geminata is a single-celled alga found in lakes, streams, and rivers. Nuisance blooms of D geminata affect the diversity, abundance, and productivity of other aquatic organisms. Because D geminata can be transported by humans on waders and other gear, accurate spatial prediction of habitat suitability is urgently needed for early detection and rapid response, as well as for evaluation of monitoring and control programs. We compared four modeling methods to predict D geminatas habitat distribution; two methods use presence–absence data (logistic regression and classification and regression tree [CART]), and two involve presence data (maximum entropy model [Maxent] and genetic algorithm for rule-set production [GARP]). Using these methods, we evaluated spatially explicit, bioclimatic and environmental variables as predictors of diatom distribution. The Maxent model provided the most accurate predictions, followed by logistic regression, CART, and GARP. The most suitable habitats we...