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

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.

/pdf/riparian-zones-as-havens-for-exotic-plant-species-in-the-5cmx26rwqv.pdf

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

Maintaining native plant diversity, detecting exotic species, and monitoring rare species are becoming important objectives in rangeland conservation. Four rangeland vegetation sampling techniques were compared to see how well they captured local plant diversity. The methods tested included the commonly used Parker transects, Daubenmire transects as modified by the USDA Forest Service, a new transect and "large quadrat" design proposed by the USDA Agricultural Research Service, and the Modified-Whittaker multi-scale vegetation plot. The 4 methods were superimposed in shortgrass steppe, mixed grass prairie, northern mixed prairie, and tallgrass prairie in the Central Grasslands of the United States with 4 replicates in each prairie type. Analysis of variance tests showed significant method effects and prairie type effects, but no significant method X type interactions for total species richness, the number of native species, the number of species with less than 1% cover, and the time required for sampling. The methods behaved similarly in each prairie type under a wide variety of grazing regimes. The Parker, large quadrat, and Daubenmire transects significantly underestimated the total species richness and the number of native species in each prairie type, and the number of species with less than 1% cover in all but the tallgrass prairie type. The transect techniques also consistently missed half the exotic species, including noxious weeds, in each prairie type. The Modified-Whittaker method, which included an exhaustive search for plant species in a 20 x 50 m plot, served as the baseline for species richness comparisons. For all prairie types, the Modified-Whittaker plot captured an average of 42.9 (+/- 2.4; 1 S.E.) plant species per site compared to 15.9 (+/- 1.3), 18.9 (+/- 1.2), and 22.8 (+/- 1.6) plant species per site using the Parker, large quadrat, and Daubenmire transect methods, respectively. The 4 methods captured most of the dominant species at each site and thus produced similar results for total foliar cover and soil cover. The detection and measurement of exotic plant species were greatly enhanced by using ten 1 m2 subplots in a multi-scale sampling design and searching a larger area (1,000 m2) at each site. Even with 4 replicate sites, the transect methods usually captured, and thus would monitor, 36 to 66% of the plant species at each site. To evaluate the status and trends of common, rare, and exotic plant species at local, regional, and national scales, innovative, multi-scale methods must replace the commonly used transect methods of the past.

/pdf/comparison-of-rangeland-vegetation-sampling-techniques-in-1whh5admlb.pdf

Kelly A. Bull

Papers

Exotic plant species invade hot spots of native plant diversity

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

Comparison of rangeland vegetation sampling techniques in the Central Grasslands