Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

Humans are altering the composition of biological communities through a variety of activities that increase rates of species invasions and species extinctions, at all scales, from local to global. These changes in components of the Earth's biodiversity cause concern for ethical and aesthetic reasons, but they also have a strong potential to alter ecosystem properties and the goods and services they provide to humanity. Ecological experiments, observations, and theoretical developments show that ecosystem properties depend greatly on biodiversity in terms of the functional characteristics of organisms present in the ecosystem and the distribution and abundance of those organisms over space and time. Species effects act in concert with the effects of climate, resource availability, and disturbance regimes in influencing ecosystem properties. Human activities can modify all of the above factors; here we focus on modification of these biotic controls. The scientific community has come to a broad consensus on many aspects of the re- lationship between biodiversity and ecosystem functioning, including many points relevant to management of ecosystems. Further progress will require integration of knowledge about biotic and abiotic controls on ecosystem properties, how ecological communities are struc- tured, and the forces driving species extinctions and invasions. To strengthen links to policy and management, we also need to integrate our ecological knowledge with understanding of the social and economic constraints of potential management practices. Understanding this complexity, while taking strong steps to minimize current losses of species, is necessary for responsible management of Earth's ecosystems and the diverse biota they contain.

/pdf/effects-of-biodiversity-on-ecosystem-functioning-a-consensus-13j8vabcus.pdf

Effects of biodiversity on ecosystem functioning: a consensus of current knowledge

Human alteration of the global environment has triggered the sixth major extinction event in the history of life and caused widespread changes in the global distribution of organisms. These changes in biodiversity alter ecosystem processes and change the resilience of ecosystems to environmental change. This has profound consequences for services that humans derive from ecosystems. The large ecological and societal consequences of changing biodiversity should be minimized to preserve options for future solutions to global environmental problems.

Consequences of changing biodiversity

Summary
1 The invasion of habitats by non-native plant and animal species is a global phenomenon with potentially grave consequences for ecological, economic, and social systems. Unfortunately, to date, the study of invasions has been primarily anecdotal and resistant to generalization.

2 Here, we use insights from experiments and from long-term monitoring studies of vegetation to propose a new theory in which fluctuation in resource availability is identified as the key factor controlling invasibility, the susceptibility of an environment to invasion by non-resident species. The theory is mechanistic and quantitative in nature leading to a variety of testable predictions.

3 We conclude that the elusive nature of the invasion process arises from the fact that it depends upon conditions of resource enrichment or release that have a variety of causes but which occur only intermittently and, to result in invasion, must coincide with availability of invading propagules.

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1046/j.1365-2745.2000.00473.x

Fluctuating resources in plant communities: a general theory of invasibility

Community ecology theory can be used to understand biological invasions by applying recent niche concepts to alien species and the communities that they invade. These ideas lead to the concept of ‘niche opportunity', which defines conditions that promote invasions in terms of resources, natural enemies, the physical environment, interactions between these factors, and the manner in which they vary in time and space. Niche opportunities vary naturally between communities but might be greatly increased by disruption of communities, especially if the original community members are less well adapted to the new conditions. Recent niche theory clarifies the prediction that low niche opportunities (invasion resistance) result from high species diversity. Conflicting empirical patterns of invasion resistance are potentially explained by covarying external factors. These various ideas derived from community ecology provide a predictive framework for invasion ecology.

/pdf/community-ecology-theory-as-a-framework-for-biological-2tep5nf3cd.pdf

Community ecology theory as a framework for biological invasions

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.

/pdf/exotic-plant-species-invade-hot-spots-of-native-plant-1594qms5ae.pdf

Exotic plant species invade hot spots of native plant diversity

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.

A Modified-Whittaker nested vegetation sampling method

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...

How grazing and soil quality affect native and exotic plant diversity in Rocky Mountain grasslands

Basic information on where nonnative plant spe- cies have successfully invaded is lacking. We assessed the vulnerability of 22 vegetation types (25 sets of four plots in nine study areas) to nonnative plant invasions in the north- central United States. In general, habitats with high native species richness were more heavily invaded than species- poor habitats, low-elevation areas were more invaded than high-elevation areas, and riparian zones were more invaded than nearby upland sites. For the 100 1000-m 2 plots (across all vegetation types), 50% of the variation in nonnative species richness was explained by longitude, latitude, native plant species richness, soil total percentage nitrogen, and mean maximum July temperature (n 100 plots; P 0.001). At the vegetation-type scale (n 25 sets of four 1000-m 2 plots/ type), 64% of the variation in nonnative species richness was explained by native plant species richness, elevation, and Oc- tober to June precipitation (P 0.001). The foliar cover of nonnative species (log) was strongly positively correlated with the nonnative species richness at the plot scale (r 0.77, P 0.001) and vegetation-type scale (r 0.83, P 0.001). We concluded that, at the vegetation-type and re- gional scales in the north- central United States, (1) vegetation types rich in native species are often highly vulnerable to inva- sion by nonnative plant species; (2) where several nonnative species become established, nonnative species cover can substantially increase; (3) the attributes that maintain high na- tive plant species richness (high light, water, nitrogen, and temperatures) also help maintain nonnative plant species rich- ness; and (4) more care must be taken to preserve native spe- cies diversity in highly vulnerable habitats. Invasive nonnative plant species are a growing con- cern in the north- central United States and elsewhere because some of them can poison livestock, clog water- ways, compete with cash crops, and degrade rangelands (Westbrooks 1999). In this paper, we operationally de- fine nonnative species as species from other continents. Managers of national parks, wildlife refuges, and other natural areas are also concerned because of the poten- tial negative effects of nonnative plant species on native

/pdf/assessing-vulnerability-to-invasion-by-nonnative-plant-kli5fsarzo.pdf

Assessing vulnerability to invasion by nonnative plant species at multiple spatial scales.

Studies to identify gaps in the protection of habitat for speciesof concern have been inconclusive and hampered by single-scale orpoor multi-scale sampling methods, large minimum mapping units(MMU's of 2 ha to 100 ha), limited and subjectively selected fieldobservations, and poor mathematical and ecological models. Weovercome these obstacles with improved multi-scale samplingtechniques, smaller MMU's (< 0.02 ha), an unbiased sampling designbased on double sampling, improved mathematical models includingspecies-area curves corrected for habitat heterogeneity, andgeographic information system-based ecological models. We applythis landscape analysis approach to address resource issues inRocky Mountain National Park, Colorado. Specifically, we quantifythe effects of elk grazing on plant diversity, identify areas ofhigh or unique plant diversity needing increased protection, andevaluate the patterns of non-native plant species on thelandscape. Double sampling techniques use satellite imagery,aerial photography, and field data to stratify homogeneous andheterogeneous units and “keystone ecosystems” (ecosystems thatcontain or support a high number of species or have distinctivespecies compositions). We show how a multi-scale vegetationsampling design, species-area curves, analyses of within- andbetween-vegetation type species overlap, and geographic informationsystem (GIS) models can be used to quantify landscape-scalepatterns of vascular plant diversity in the Park. The new multi-scale vegetation plot techniques quickly differentiated plantspecies differences in paired study sites. Three plots in the OuzelBurn area (burned in 1978) contained 75 plant species, while only17 plant species were found in paired plots outside the burn.Riparian areas contained 109 plant species, compared to just 55species in paired plots in adjacent forests. However, plant speciesrichness patterns inside and outside elk exclosures were morecomplex. One elk exclosure contained more species than its adjacentopen range (52 species inside and 48 species outside). Two elkexclosures contained fewer species inside than outside (105 and 41species inside and 112 and 74 species outside, respectively).However, there was only 26% to 48% overlap (using Jaccard'sCoefficient) of plant species composition inside and outside theexclosures. One elk exclosure had 13% cover of non-indigenousspecies inside the exclosure compared to 4% outside, butnon-indigenous species cover varied by location. We compared plantdiversity patterns from vegetation maps made with 100 ha, 50 ha, 2ha, and 0.02 ha MMU's in the 754 ha Beaver Meadows study area usingfour 0.025 ha and twenty-one 0.1 ha multi-scale vegetation plots.Preliminary data suggested that the 2 ha MMU provided an accurateestimate of the number of plant species (–14%) for a study area,but the number of habitats (polygons) was reduced by 67%, andaspen, a unique and important habitat type, was missed entirely. Wedescribe a hypothesis-driven approach to the design andimplementation of geospatial databases for local resourcemonitoring and ecosystem management.

Lisa D. Schell

Papers

Exotic plant species invade hot spots of native plant diversity

A Modified-Whittaker nested vegetation sampling method

How grazing and soil quality affect native and exotic plant diversity in Rocky Mountain grasslands

Assessing vulnerability to invasion by nonnative plant species at multiple spatial scales.

Landscape analysis of plant diversity