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

An essential textbook for any student or researcher in biology needing to design experiments, sample programs or analyse the resulting data The text begins with a revision of estimation and hypothesis testing methods, covering both classical and Bayesian philosophies, before advancing to the analysis of linear and generalized linear models Topics covered include linear and logistic regression, simple and complex ANOVA models (for factorial, nested, block, split-plot and repeated measures and covariance designs), and log-linear models Multivariate techniques, including classification and ordination, are then introduced Special emphasis is placed on checking assumptions, exploratory data analysis and presentation of results The main analyses are illustrated with many examples from published papers and there is an extensive reference list to both the statistical and biological literature The book is supported by a website that provides all data sets, questions for each chapter and links to software

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Experimental Design and Data Analysis for Biologists

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.

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

Collinearity refers to the non independence of predictor variables, usually in a regression-type analysis. It is a common feature of any descriptive ecological data set and can be a problem for parameter estimation because it inflates the variance of regression parameters and hence potentially leads to the wrong identification of relevant predictors in a statistical model. Collinearity is a severe problem when a model is trained on data from one region or time, and predicted to another with a different or unknown structure of collinearity. To demonstrate the reach of the problem of collinearity in ecology, we show how relationships among predictors differ between biomes, change over spatial scales and through time. Across disciplines, different approaches to addressing collinearity problems have been developed, ranging from clustering of predictors, threshold-based pre-selection, through latent variable methods, to shrinkage and regularisation. Using simulated data with five predictor-response relationships of increasing complexity and eight levels of collinearity we compared ways to address collinearity with standard multiple regression and machine-learning approaches. We assessed the performance of each approach by testing its impact on prediction to new data. In the extreme, we tested whether the methods were able to identify the true underlying relationship in a training dataset with strong collinearity by evaluating its performance on a test dataset without any collinearity. We found that methods specifically designed for collinearity, such as latent variable methods and tree based models, did not outperform the traditional GLM and threshold-based pre-selection. Our results highlight the value of GLM in combination with penalised methods (particularly ridge) and threshold-based pre-selection when omitted variables are considered in the final interpretation. However, all approaches tested yielded degraded predictions under change in collinearity structure and the ‘folk lore’-thresholds of correlation coefficients between predictor variables of |r| >0.7 was an appropriate indicator for when collinearity begins to severely distort model estimation and subsequent prediction. The use of ecological understanding of the system in pre-analysis variable selection and the choice of the least sensitive statistical approaches reduce the problems of collinearity, but cannot ultimately solve them.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1600-0587.2012.07348.x

Collinearity: a review of methods to deal with it and a simulation study evaluating their performance

▪ Abstract The focus of most ideas on diversity maintenance is species coexistence, which may be stable or unstable. Stable coexistence can be quantified by the long-term rates at which community members recover from low density. Quantification shows that coexistence mechanisms function in two major ways: They may be (a) equalizing because they tend to minimize average fitness differences between species, or (b) stabilizing because they tend to increase negative intraspecific interactions relative to negative interspecific interactions. Stabilizing mechanisms are essential for species coexistence and include traditional mechanisms such as resource partitioning and frequency-dependent predation, as well as mechanisms that depend on fluctuations in population densities and environmental factors in space and time. Equalizing mechanisms contribute to stable coexistence because they reduce large average fitness inequalities which might negate the effects of stabilizing mechanisms. Models of unstable coexitence...

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Mechanisms of Maintenance of Species Diversity

An oligohaline tidal marsh on the northern shore of Lake Pontchartrain, LA was characterized with respect to the distributions and abundances of plant species over spatial and temporal gradients using Detrended Correspondence Analysis (DCA). In addition, the species distributions were correlated to several physical environmental factors using Detrended Canonical Correspondence Analysis (DCCA). The distributions of species were best correlated with distance from Lake Pontchartrain, and to a lesser extent with elevation and substrate organic matter. They were least correlated with mean soil salinity (referred to here as background salinity). Of the three mid-seasonal dominant species, the perennial grass, Spartina patens, is the most salt tolerant and was found closest to the lake. Further inland the dominant perennial was Sagittaria lancifolia, which has a salt tolerance less than that of Spartina patens. The perennial sedge, Cladium jamaicense, which is the least salt tolerant of the three, was dominant furthest inland. Background salinity levels were generally low (<5 ppt.) and did not explain species distributions. We hypothesize that the distribution of species is regulated by occasional storm-generated ‘salt pulses’ that generate strong, short-lived salinity gradients as a function of distance from the lake. Biotic interactions likely also play a role in structuring the plant community. The distributions of several annuals depended on the size and life history of the mid-seasonal dominant perennials. Most of the annuals frequently co-occurred with Sagittaria lancifolia, which was the shortest in stature and had the least persistent canopy of the three mid-seasonal dominant perennials.

Plant community structure in an oligohaline tidal marsh.

1 – Perspectives on Plant Competition: Some Introductory Remarks

The objective of this study was to experimentally evaluate the effects of simulated herbivory on the ability of a freshwater marsh plant to recover from temporary saltwater intrusion such as can be caused by tropical storms. Sods containingSagittaria lancifolia, a dominant plant in interior coastal marshes, were manipulated in the field so as to subject plants to a pulse of 15‰ salt water for a duration of 1 wk. In addition to the exposure to salt water, some plants were also subjected to both short-term and long-term flooding treatments of 20 cm, and to simulated herbivory (clipping). Following exposure to salt water, plants were allowed to recover over the winter and were harvested the next June. Neither simulated herbivory, nor salinity, nor flooding caused any long-term effect either singly or in pairwise combinations. However, when plants were subjected to herbivory, salt water, and flooding simultaneously, reduced growth and plant death occurred. These results suggest that high levels of grazing by herbivores may increase the susceptibility of coastal marsh plants to damage from saltwater intrusion. *** DIRECT SUPPORT *** A01BY073 00002

The potential impact of herbivores on the susceptibility of the marsh plant Sagittaria lancifolia to saltwater intrusion in coastal wetlands

Cogongrass is a nonindigenous species perceived to threaten native communities of the southeastern United States through modification of species composition and alteration of community processes. To examine how gap size and disturbance type influence the invasion of wet pine savannas by cogongrass, we performed three field experiments to evaluate the response of cogongrass seeds and transplanted seedlings to four different gap sizes, four types of site disturbance, and recent burning of savanna vegetation. Cogongrass germinated, survived, and grew in all gap sizes, from 0 to 100 cm in diameter. Similarly, disturbance type had no effect on germination or seedling and transplant survival. Tilling, however, significantly enhanced transplanted seedling growth, resulting in a tenfold increase in biomass over the other disturbance types. Seedling survival to 1 and 2 mo was greater in burned savanna than unburned savanna, although transplant survival and growth were not affected by burning. Results of this study suggest that cogongrass can germinate, survive, and grow in wet pine savanna communities regardless of gap size or type of disturbance, including burning. Burning of savanna vegetation may enhance establishment by improving early seedling survival, and soil disturbance can facilitate invasion of cogongrass by enhancing plant growth.

https://bsapubs.onlinelibrary.wiley.com/doi/pdf/10.2307/2656721

The effects of gap size and disturbance type on invasion of wet pine savanna by cogongrass, Imperata cylindrica (Poaceae).

This paper tests the hypothesis that plant size can affect a species' relative competitive ability; i.e., species that are good competitors as adults may not also be good competitors as juveniles. Seeds of two Typha species were sown along a miniature gradient in water depth, both in monocultures and in mixtures, to ascertain the relative competitive abilities ofjuveniles. In monoculture, T. latifolia was found to germinate and grow better at slightly higher elevations than T. domingensis. In mixture, T. latifolia was the better competitor above the water table while T. domingensis dominated below the water table. Summed over the entire gradient, the two species did not differ significantly in their overall competitive abilities despite the fact that T. latifolia has larger seeds. Adult competitive abilities were studied experimentally in a large artificial pond by using transplants planted at low initial densities. When each species was grown in monoculture, T. latifolia grew best in shallow water with little growth in water -1.0 m deep, and T. domingensis had peak shoot growth in deep water (0.8-0.9 m) with comparatively less growth in shallow water. When grown in mixture, T. latifolia was the dominant competitor over virtually the entire gradient. Therefore, I conclude that relative competitive ability in Typha is size dependent. The factors most likely to cause the observed shift in relative competitive abilities are (1) the onset of flowering, (2) changes in morphology and physiology associated with growth and development, and (3) changes in the factors limiting growth as plants increase in size. Regardless of the cause, the observed relationship between size and competitive ability suggests that the type of initial propagules used in competition experiments can affect the outcome, and that in some cases, the outcome of competition may be dependent upon the initial density of propagules.

James B. Grace

Papers

Plant community structure in an oligohaline tidal marsh.

1 – Perspectives on Plant Competition: Some Introductory Remarks

The potential impact of herbivores on the susceptibility of the marsh plant Sagittaria lancifolia to saltwater intrusion in coastal wetlands

The effects of gap size and disturbance type on invasion of wet pine savanna by cogongrass, Imperata cylindrica (Poaceae).

Juveniles vs. Adult Competitive Abilities in Plants: Size‐dependence in Cattails (Typha)