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

Pseudoreplication is defined. as the use of inferential statistics to test for treatment effects with data from experiments where either treatments are not replicated (though samples may be) or replicates are not statistically independent. In ANOVA terminology, it is the testing for treatment effects with an error term inappropriate to the hypothesis being considered. Scrutiny of 176 experi- mental studies published between 1960 and the present revealed that pseudoreplication occurred in 27% of them, or 48% of all such studies that applied inferential statistics. The incidence of pseudo- replication is especially high in studies of marine benthos and small mammals. The critical features of controlled experimentation are reviewed. Nondemonic intrusion is defined as the impingement of chance events on an experiment in progress. As a safeguard against both it and preexisting gradients, interspersion of treatments is argued to be an obligatory feature of good design. Especially in small experiments, adequate interspersion can sometimes be assured only by dispensing with strict random- ization procedures. Comprehension of this conflict between interspersion and randomization is aided by distinguishing pre-layout (or conventional) and layout-specifit alpha (probability of type I error). Suggestions are offered to statisticians and editors of ecological j oumals as to how ecologists' under- standing of experimental design and statistics might be improved.

/pdf/pseudoreplication-and-the-design-of-ecological-field-44o8kzdaoi.pdf

Pseudoreplication and the Design of Ecological Field Experiments

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

Species richness is a fundamental measurement of community and regional diversity, and it underlies many ecological models and conservation strategies. In spite of its importance, ecologists have not always appreciated the effects of abundance and sampling effort on richness measures and comparisons. We survey a series of common pitfalls in quantifying and comparing taxon richness. These pitfalls can be largely avoided by using accumulation and rarefaction curves, which may be based on either individuals or samples. These taxon sampling curves contain the basic information for valid richness comparisons, including category‐subcategory ratios (species-to-genus and species-toindividual ratios). Rarefaction methods ‐ both sample-based and individual-based ‐ allow for meaningful standardization and comparison of datasets. Standardizing data sets by area or sampling effort may produce very different results compared to standardizing by number of individuals collected, and it is not always clear which measure of diversity is more appropriate. Asymptotic richness estimators provide lower-bound estimates for taxon-rich groups such as tropical arthropods, in which observed richness rarely reaches an asymptote, despite intensive sampling. Recent examples of diversity studies of tropical trees, stream invertebrates, and herbaceous plants emphasize the importance of carefully quantifying species richness using taxon sampling curves.

/pdf/quantifying-biodiversity-procedures-and-pitfalls-in-the-2ut2f3nw4g.pdf

Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness

A strategy to understand the microbial components of the human genetic and metabolic landscape and how they contribute to normal physiology and predisposition to disease.

/pdf/the-human-microbiome-project-1armeyuv2c.pdf

The human microbiome project.

How are diversity and productivity related

Summary 1 We develop and test a game-theoretic model for considering the effects of intra- and interplant competition on root proliferation and reproductive yield. 2 We predict that if space and resources per individual are held constant, plants should produce more roots per individual and less reproductive yield per individual as the number of plants sharing the combined space increases. 3 We tested the predictions using soybean plants ( Glycine max ) cultivated in the glasshouse either as owners or as two individuals sharing twice the space and nutrients. 4 Sharing individuals produced 85% more root mass than owners. Owners, however, produced 30% more reproductive yield per plant (dry mass of seeds), as a result of significantly more seed pods (8.70 vs. 7.66), more seeds per pod (1.87 vs. 1.72) and larger seeds (0.205 vs. 0.195 g seed ‐1 ), than did sharing individuals. 5 Total plant biomass did not differ between owners and sharing individuals, but owners had significantly higher shoot to root ratios, produced significantly more seeds per unit root mass, and allocated a significantly higher percentage of total biomass production to seeds. 6 Possession of an evolutionarily stable strategy (ESS) of root competition suggests that different roots and parts of a plant assess and respond to opportunities in a manner that maximizes the good of the whole plant. Thus, plants may be more sophisticated and share more in common with animals in their non-cognitive behaviours than previously thought. A plant operating as a co-ordinated whole should, all else being equal, first proliferate roots in unoccupied soil, then in soil occupied by a conspecific competitor, and lastly in soil already occupied by its own roots.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1046/j.0022-0477.2001.00609.x

Tragedy of the commons as a result of root competition

We have tested two hypotheses which may explain two different patterns which underlie coexistence in two species of desert gerbils (Gerbillus allenbyi and G. pyramidum). The two patterns are temporal partitioning of foraging activity and shared preference habitat selection. When sympatric, G. pyramidum uses the early part of the night most heavily while G. allenbyi is most active in the late part of the night. Although both species prefer the same habitat type (semistabilized sand dune), in the presence of G, pyramidum, G. allenbyi utilizes mainly the secondary habitat type (stabilized sand dune)

Interference competition and temporal and habitat partitioning in two gerbil species

Tilman has developed a model to predict the number of plant species that can coexist competitively on a limited resource base. Species diversity first increases over low resource supplies, then declines as the environment becomes richer. Although Tilman 's model was developed to describe interspecific interactions between plant species, it may also apply to animal species. Tilman questions whether animals specialize on particular proportions of nutrients. However, we believe animals probably specialize on relatively subtle microhabitat differences, especially in a multispecies competitive regime. Thus, microhabitats may act like nutrients. We hypothesize that animal species, too, show a peaked curve of diversity over productivity. The present data provide a confirmation of the hypothesis using rodent species. We have investigated the number of rodent species along a geographical gradient of increasing rainfall. The gradient extends from extremely poor desert habitats to those with annual rainfall over 300 mm. Because of the aridity , precipitation reflects productivity. The diversity pattern in desert rodents agrees with that predicted by Tilman for plants. It even possesses similar asymmetry, rising steeply then falling slowly. The pattern is duplicated in rocky and sandy habitats, each of which has a distinct and almost nonoverlapping assemblage of species. As mean precipitation is closely correlated with the variability of precipitation, the diversity pattern might also be caused by a decline in the frequency of disturbances, models for which have been proposed by several investigators.

Tilman's predicted productivity–diversity relationship shown by desert rodents

Predation plays an important role in ecological communities by affecting prey behavior such as foraging and by physical removal of individual prey. In regard to foraging, animals such as desert rodents often balance conflicting demands for food and safety. This has been studied in the field by indirectly manipulating predatory risk through the alteration of cues associated with increased risk such as cover or illumination. It has also been studied by directly manipulating the presence of predators in aviaries. Here, we report on experiments in which we directly manipulated actual predatory risk to desert rodents in the field. We conducted a series of experiments in the field using a trained barn owl (Tyto alba) to investigate how two species of coexisting gerbils (Gerbillus allenbyi and G. pyramidum) respond to various cues of predatory risk in their natural environment. The gerbils responded to risk of predation, in the form of owl flights and owl hunger calls, by reducing their activity in the risky plot relative to the control plot. The strongest response was to owl flights and the weakest to recorded hunger calls of owls. Furthermore, when risk of predation was relatively high, as in the case with barn owl flights, both gerbil species mostly limited their activity to the safer bush microhabitat. The response of the gerbils to risk of predation disappeared very quickly following removal of the treatment, and the gerbils returned to normal levels of activity within the same night. The gerbils did not respond to experimental cues (alarm clock), the presence of the investigators, the presence of a quiet owl, and recorded “white noise”. Using trained barn owls, we were able to effectively manipulate actual risk of predation to gerbils in natural habitats and to quantify how gerbils alter their behavior in order to balance conflicting demands of food and safety. The method allows assessment of aspects of behavior, population interactions, and community characteristics involving predation in natural habitats.

Zvika Abramsky

Papers

How are diversity and productivity related

Tragedy of the commons as a result of root competition

Interference competition and temporal and habitat partitioning in two gerbil species

Tilman's predicted productivity–diversity relationship shown by desert rodents

The effect of barn owls (Tyto alba) on the activity and microhabitat selection of Gerbillus allenbyi and G. pyramidum.