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

Biotic invaders are species that establish a new range in which they proliferate, spread, and persist to the detriment of the environment. They are the most important ecological outcomes from the unprecedented alterations in the distribution of the earth's biota brought about largely through human transport and commerce. In a world without borders, few if any areas remain sheltered from these im- migrations. The fate of immigrants is decidedly mixed. Few survive the hazards of chronic and stochastic forces, and only a small fraction become naturalized. In turn, some naturalized species do become invasive. There are several potential reasons why some immigrant species prosper: some escape from the constraints of their native predators or parasites; others are aided by human-caused disturbance that disrupts native communities. Ironically, many biotic invasions are apparently facilitated by cultivation and husbandry, unintentional actions that foster immigrant populations until they are self-perpetuating and uncontrollable. Whatever the cause, biotic invaders can in many cases inflict enormous environmental damage: (1) Animal invaders can cause extinctions of vulnerable native species through predation, grazing, competition, and habitat alteration. (2) Plant invaders can completely alter the fire regime, nutrient cycling, hydrology, and energy budgets in a native ecosystem and can greatly diminish the abundance or survival of native species. (3) In agriculture, the principal pests of temperate crops are nonindigenous, and the combined expenses of pest control and crop losses constitute an onerous "tax" on food, fiber, and forage production. (4) The global cost of virulent plant and animal diseases caused by parasites transported to new ranges and presented with susceptible new hosts is currently incalculable. Identifying future invaders and taking effective steps to prevent their dispersal and establishment con- stitutes an enormous challenge to both conservation and international commerce. Detection and management when exclusion fails have proved daunting for varied reasons: (1) Efforts to identify general attributes of future invaders have often been inconclusive. (2) Predicting susceptible locales for future invasions seems even more problematic, given the enormous differences in the rates of arrival among potential invaders. (3) Eradication of an established invader is rare, and control efforts vary enormously in their efficacy. Successful control, however, depends more on commitment and continuing diligence than on the efficacy of specific tools themselves. (4) Control of biotic invasions is most effective when it employs a long-term, ecosystem- wide strategy rather than a tactical approach focused on battling individual invaders. (5) Prevention of invasions is much less costly than post-entry control. Revamping national and international quarantine laws by adopting a "guilty until proven innocent" approach would be a productive first step. Failure to address the issue of biotic invasions could effectively result in severe global consequences, including wholesale loss of agricultural, forestry, and fishery resources in some regions, disruption of the ecological processes that supply natural services on which human enterprise depends, and the creation of homogeneous, impoverished ecosystems composed of cosmopolitan species. Given their current scale, biotic invasions have taken their place alongside human-driven atmospheric and oceanic alterations as major agents of global change. Left unchecked, they will influence these other forces in profound but still unpredictable ways.

/pdf/biotic-invasions-causes-epidemiology-global-consequences-and-1uc39f4dh2.pdf

Biotic invasions: causes, epidemiology, global consequences, and control

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

All terrestrial ecosystems consist of aboveground and belowground components that interact to influence community- and ecosystem-level processes and properties. Here we show how these components are closely interlinked at the community level, reinforced by a greater degree of specificity between plants and soil organisms than has been previously supposed. As such, aboveground and belowground communities can be powerful mutual drivers, with both positive and negative feedbacks. A combined aboveground-belowground approach to community and ecosystem ecology is enhancing our understanding of the regulation and functional significance of biodiversity and of the environmental impacts of human-induced global change phenomena.

http://science.sciencemag.org/content/sci/304/5677/1629.full.pdf

Ecological linkages between aboveground and belowground biota.

Interactions among organisms take place within a complex milieu of abiotic and biotic processes, but we generally study them as solitary phenomena. Complex combinations of negative and positive interactions have been identified in a number of plant communities. The importance of these two processes in structuring plant communities can best be understood by comparing them along gradients of abiotic stress, consumer pressure, and among different life stages, sizes, and densities of the interacting species. Here, we discuss the roles of life stage, physiology, indirect interactions, and the physical environment on the balance of competition and facilitation in plant communities.

/pdf/competition-and-facilitation-a-synthetic-approach-to-51azsyo86c.pdf

Competition and facilitation: a synthetic approach to interactions in plant communities

Myrica faya, an introduced actinorhizal nitrogen fixer, in invading young volcanic sites in Hawaii Volcanoes National Park. We examined the population biology of the invader and ecosystem-level consequences of its invasion in open-canopied forests resulting from volcanic cinder-fall. Although Myrica faya is nominally dioecious, both males and females produce large amounts of fruit that are utilized by a number of exotic and native birds, particularly the exotic Zosterops japonica. In areas of active colonization, Myrica seed rain under perch trees of the dominant native Metrosideros polymorpha ranged from 6 to 60 seeds m{sup {minus}2} yr{sup {minus}1}; no seeds were captured in the open. Planted seeds of Myrica also germinated an established better under isolated individuals of Metrosideros than in the open. Diameter growth of Myrica is > 15-fold greater than that of Metrosideros, and the Myrica population is increasing rapidly. Rates of nitrogen fixation were measured using the acetylene reduction assay calibrated with {sup 15}N. Myrica nodules reduced acetylene at between 5 and 20 {mu}mol g{sup {minus}1} h{sup {minus}1}, a rate that extrapolated to nitrogen fixation of 18 kg ha{sup {minus}1} in a densely colonized site. By comparison, all native sources of nitrogen fixation summed to 0.2 kg ha{sup {minus}1}more » yr{sup {minus}1}, and precipitation added < 4 kg ha{sup {minus}1} yr{sup {minus}1}. Measurements of litter decomposition and nitrogen release, soil nitrogen mineralization, and plant growth in bioassays all demonstrated that nitrogen fixed by Myrica becomes available to other organisms as well. We conclude that biological invasion by Myrica faya alters ecosystem-level properties in this young volcanic area; at least in this case, the demography and physiology of one species controls characteristics of a whole ecosystem.« less

Biological invasion by Myrica faya in Hawai'i: plant demography, nitrogen fixation, ecosystem effects

In primary succession following deglaciation at Glacier Bay, Alaska, we tested the hypothesis that the major effect of initial nitrogen-fixing colonizers is to facilitate establishment of late-successional dom- inants and that other possible causes of successional change (e.g., life history factors governing seed rain and competitive interactions among species) need not be invoked. Environment changed dramatically through the first 200 yr of succession. Soil organic matter increased 10-fold in the upper mineral soil with corresponding increases in soil moisture, total nitrogen (N), and capacity to support plant growth and declines in bulk density, pH, and total phosphorus (P). Plant growth in pioneer soils tended to be simultaneously limited by both N and P, as well as by unknown factors (perhaps lack of mycorrhizae), whereas only P limited growth in older soils. Light availability to seedlings declined through succession. Early-successional species (Epilobium latifolium, Dryas drummondii) had smaller seeds, younger age at first reproduction, shorter life-span, and shorter height at maturity than did mid-successional (alder, Alnus sinuata) and late-successional species (sitka spruce, Picea sitchensis). Seed rain of alder and spruce was negligible in the pioneer stage, increased prior to the stage in which a species was dominant, and was greatest in the stage in which a species dominated. Vegetation in each successional stage inhibited germination and initial establishment of sown alder and spruce seeds (except a tendency of the "black-crust" algal/microbial community in the pioneer stage to enhance survivorship). Removal of the surface litter layer generally enhanced germination and survi- vorship, particularly of alder. Comparisons of germination in the greenhouse and the field indicated that climatic or indirect vegetation effects (e.g., differential seed predation) and allelopathy also reduced germination and establishment in vegetated communities. Naturally occurring spruce seedlings grew most rapidly in the Dryas and alder stages and most slowly in the spruce stage. Similarly, growth of spruce seedlings transplanted into each successional stage was facilitated by the Dryas (nonsignificantly) and alder stages but inhibited by the spruce stage, relative to earlier successional stages. Facilitation of growth of natural and transplanted spruce seedlings by Dryas and alder stages was associated with higher N and P uptake and tissue nutrient concentrations, whereas nutrient uptake and concentration in spruce seedlings declined in the spruce stage. By contrast, transplanted alder seedlings grew rapidly and accu- mulated most nutrients in the pioneer stage and were strongly inhibited by subsequent stages. The facilitative effect of Dryas and alder comes primarily from inputs of organic matter and associated N. Addition of alder litter stimulated nutrient uptake and growth of transplanted spruce seedlings in the pioneer and Dryas stages, whereas shading had no effect on growth of spruce seedlings. Root trenching and planting of spruce near isolated alders indicated that, although the net effect of alder is facilitative, alder also inhibits growth of spruce seedlings through competition for soil resources. Strong root competition also occurs in the spruce stage. Alder competitively inhibits Dryas, primarily by shading but also through the physical and allelopathic effects of its litter. In general, both at Glacier Bay and elsewhere, life history traits determine the pattern of succession. Changes in competitive balance accompanying successional changes in environment provide the mechanism for changes in species dominance. Initial site conditions (and facilitation, where present) influence the rate of change and final state of community composition and productivity. We conclude that no single factor or mechanism fully accounts for primary succession at Glacier Bay.

https://www.jstor.org/stable/pdfplus/2937039.pdf

Mechanisms of Primary Succession Following Deglaciation at Glacier Bay, Alaska

Summary 1. Chronosequences and associated space-for-time substitutions are an important and often necessary tool for studying temporal dynamics of plant communities and soil development across multiple time-scales. However, they are often used inappropriately, leading to false conclusions about ecological patterns and processes, which has prompted recent strong criticism of the approach. Here, we evaluate when chronosequences may or may not be appropriate for studying community and ecosystem development. 2. Chronosequences are appropriate to study plant succession at decadal to millennial time-scales when there is evidence that sites of different ages are following the same trajectory. They can also be reliably used to study aspects of soil development that occur between temporally linked sites over time-scales of centuries to millennia, sometimes independently of their application to shorter-term plant and soil biological communities. 3. Some characteristics of changing plant and soil biological communities (e.g. species richness, plant cover, vegetation structure, soil organic matter accumulation) are more likely to be related in a predictable and temporally linear manner than are other characteristics (e.g. species composition and abundance) and are therefore more reliably studied using a chronosequence approach. 4. Chronosequences are most appropriate for studying communities that are following convergent successional trajectories and have low biodiversity, rapid species turnover and low frequency and severity of disturbance. Chronosequences are least suitable for studying successional trajectories that are divergent, species-rich, highly disturbed or arrested in time because then there are often major difficulties in determining temporal linkages between stages. 5. Synthesis. We conclude that, when successional trajectories exceed the life span of investigators and the experimental and observational studies that they perform, temporal change can be successfully explored through the judicious use of chronosequences.

/pdf/the-use-of-chronosequences-in-studies-of-ecological-1c5s5077we.pdf

The use of chronosequences in studies of ecological succession and soil development

During succession, ecosystem development occurs; but in the long-term absence of catastrophic disturbance, a decline phase eventually follows. We studied six long-term chronosequences, in Australia, Sweden, Alaska, Hawaii, and New Zealand; for each, the decline phase was associated with a reduction in tree basal area and an increase in the substrate nitrogen-to-phosphorus ratio, indicating increasing phosphorus limitation over time. These changes were often associated with reductions in litter decomposition rates, phosphorus release from litter, and biomass and activity of decomposer microbes. Our findings suggest that the maximal biomass phase reached during succession cannot be maintained in the long-term absence of major disturbance, and that similar patterns of decline occur in forested ecosystems spanning the tropical, temperate, and boreal zones.

https://science.sciencemag.org/content/sci/305/5683/509.full.pdf

Lawrence R. Walker

Papers

Competition and facilitation: a synthetic approach to interactions in plant communities

Biological invasion by Myrica faya in Hawai'i: plant demography, nitrogen fixation, ecosystem effects

Mechanisms of Primary Succession Following Deglaciation at Glacier Bay, Alaska

The use of chronosequences in studies of ecological succession and soil development

Ecosystem Properties and Forest Decline in Contrasting Long-Term Chronosequences