In the early 1980s, a strategy for graphical representation of multivariate (multi-species) abundance data was introduced into marine ecology by, among others, Field, et al. (1982). A decade on, it is instructive to: (i) identify which elements of this often-quoted strategy have proved most useful in practical assessment of community change resulting from pollution impact; and (ii) ask to what extent evolution of techniques in the intervening years has added self-consistency and comprehensiveness to the approach. The pivotal concept has proved to be that of a biologically-relevant definition of similarity of two samples, and its utilization mainly in simple rank form, for example ‘sample A is more similar to sample B than it is to sample C’. Statistical assumptions about the data are thus minimized and the resulting non-parametric techniques will be of very general applicability. From such a starting point, a unified framework needs to encompass: (i) the display of community patterns through clustering and ordination of samples; (ii) identification of species principally responsible for determining sample groupings; (iii) statistical tests for differences in space and time (multivariate analogues of analysis of variance, based on rank similarities); and (iv) the linking of community differences to patterns in the physical and chemical environment (the latter also dictated by rank similarities between samples). Techniques are described that bring such a framework into place, and areas in which problems remain are identified. Accumulated practical experience with these methods is discussed, in particular applications to marine benthos, and it is concluded that they have much to offer practitioners of environmental impact studies on communities.

Non-parametric multivariate analyses of changes in community structure

Hypothesis-testing methods for multivariate data are needed to make rigorous probability statements about the effects of factors and their interactions in experiments. Analysis of variance is particularly powerful for the analysis of univariate data. The traditional multivariate analogues, however, are too stringent in their assumptions for most ecological multivariate data sets. Non-parametric methods, based on permutation tests, are preferable. This paper describes a new non-parametric method for multivariate analysis of variance, after McArdle and Anderson (in press). It is given here, with several applications in ecology, to provide an alternative and perhaps more intuitive formulation for ANOVA (based on sums of squared distances) to complement the description pro- vided by McArdle and Anderson (in press) for the analysis of any linear model. It is an improvement on previous non-parametric methods because it allows a direct additive partitioning of variation for complex models. It does this while maintaining the flexibility and lack of formal assumptions of other non-parametric methods. The test- statistic is a multivariate analogue to Fisher's F-ratio and is calculated directly from any symmetric distance or dissimilarity matrix. P-values are then obtained using permutations. Some examples of the method are given for tests involving several factors, including factorial and hierarchical (nested) designs and tests of interactions.

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A new method for non-parametric multivariate analysis of variance

Background The analysis of microbial communities through DNA sequencing brings many challenges: the integration of different types of data with methods from ecology, genetics, phylogenetics, multivariate statistics, visualization and testing. With the increased breadth of experimental designs now being pursued, project-specific statistical analyses are often needed, and these analyses are often difficult (or impossible) for peer researchers to independently reproduce. The vast majority of the requisite tools for performing these analyses reproducibly are already implemented in R and its extensions (packages), but with limited support for high throughput microbiome census data. Results Here we describe a software project, phyloseq, dedicated to the object-oriented representation and analysis of microbiome census data in R. It supports importing data from a variety of common formats, as well as many analysis techniques. These include calibration, filtering, subsetting, agglomeration, multi-table comparisons, diversity analysis, parallelized Fast UniFrac, ordination methods, and production of publication-quality graphics; all in a manner that is easy to document, share, and modify. We show how to apply functions from other R packages to phyloseq-represented data, illustrating the availability of a large number of open source analysis techniques. We discuss the use of phyloseq with tools for reproducible research, a practice common in other fields but still rare in the analysis of highly parallel microbiome census data. We have made available all of the materials necessary to completely reproduce the analysis and figures included in this article, an example of best practices for reproducible research. Conclusions The phyloseq project for R is a new open-source software package, freely available on the web from both GitHub and Bioconductor.

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phyloseq: an R package for reproducible interactive analysis and graphics of microbiome census data.

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

This paper presents a new and simple method to find indicator species and species assemblages characterizing groups of sites The novelty of our approach lies in the way we combine a species relative abundance with its relative frequency of occurrence in the various groups of sites This index is maximum when all individuals of a species are found in a single group of sites and when the species occurs in all sites of that group; it is a symmetric indicator The statistical significance of the species indicator values is evaluated using a randomization procedure Contrary to TWINSPAN, our indicator index for a given species is independent of the other species relative abundances, and there is no need to use pseudospecies The new method identifies indicator species for typologies of species releves obtained by any hierarchical or nonhierarchical classification procedure; its use is independent of the classification method Because indicator species give ecological meaning to groups of sites, this method provides criteria to compare typologies, to identify where to stop dividing clusters into subsets, and to point out the main levels in a hierarchical classification of sites Species can be grouped on the basis of their indicator values for each clustering level, the heterogeneous nature of species assemblages observed in any one site being well preserved Such assemblages are usually a mixture of eurytopic (higher level) and stenotopic species (characteristic of lower level clusters) The species assemblage approach demonstrates the importance of the ''sampled patch size,'' ie, the diversity of sampled ecological combinations, when we compare the frequencies of core and satellite species A new way to present species-site tables, accounting for the hierarchical relationships among species, is proposed A large data set of carabid beetle distributions in open habitats of Belgium is used as a case study to illustrate the new method

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Species assemblages and indicator species:the need for a flexible asymmetrical approach

The robustness of quantitative measures of compositional dissimilarity between sites was evaluated using extensive computer simulations of species’ abundance patterns over one and two dimensional configurations of sample sites in ecological space. Robustness was equated with the strength, over a range of models, of the linear and monotonic (rank-order) relationship between the compositional dissimilarities and the corresponding Euclidean distances between sites measured in the ecological space. The range of models reflected different assumptions about species’ response curve shape, sampling pattern of sites, noise level of the data, species’ interactions, trends in total site abundance, and beta diversity of gradients.

Compositional dissimilarity as a robust measure of ecological distance

Summary 1. Owing to the detrimental impacts of invasive alien species, their control is often a priority for conservation management. Whereas the potential for unforeseen consequences of management is recognized, their associated complexity and costs are less widely appreciated. 2. We demonstrate that theoretically plausible trophic cascades associated with invasive species removal not only take place in reality, but can also result in rapid and drastic landscape-wide changes to ecosystems. 3. Using a combination of population data from of an invasive herbivore, plot-scale vegetation analyses, and satellite imagery, we show how a management intervention to eradicate a mesopredator has inadvertently and rapidly precipitated landscape-wide change on sub-Antarctic Macquarie Island. This happened despite the eradication being positioned within an integrated pest management framework. Following eradication of cats Felis catus in 2001, rabbit Oryctolagus cuniculus numbers increased substantially although a control action was in place ( Myxoma virus), resulting in island-wide ecosystem effects. 4. Synthesis and applications . Our results highlight an important lesson for conservation agencies working to eradicate invasive species globally; that is, risk assessment of management interventions must explicitly consider and plan for their indirect effects, or face substantial subsequent costs. On Macquarie Island, the cost of further conservation action will exceed AU$24 million.

Indirect effects of invasive species removal devastate World Heritage Island

Evaluation of statistical models used for predicting plant species distributions: Role of artificial data and theory

We compare three common types of clustering algorithms for use with community data. TWINSPAN is divi- sive hierarchical, flexible-UPGMA is agglomerative and hierarchical, and ALOC is non-hierarchical. A balanced de- sign six-factor model was used to generate 480 data sets of known characteristics. Recovery of the embedded clusters suggests that both flexible UPGMA and ALOC are signifi- cantly better than TWINSPAN. No significant difference existed between flexible UPGMA and ALOC.

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Lee Belbin

Papers

Compositional dissimilarity as a robust measure of ecological distance

Indirect effects of invasive species removal devastate World Heritage Island

Evaluation of statistical models used for predicting plant species distributions: Role of artificial data and theory

Comparing three classification strategies for use in ecology

Environmental representativeness: Regional partitioning and reserve selection