Robert P. Freckleton

Phylogenetic Analysis and Comparative Data: A Test and Review of Evidence

The question is often raised whether it is statistically necessary to control for phylogenetic associations in comparative studies. To investigate this question, we explore the use of a measure of phylogenetic correlation, λ, introduced by Pagel (1999), that normally varies between 0 (phylogenetic independence) and 1 (species’ traits covary in direct proportion to their shared evolutionary history). Simulations show λ to be a statistically powerful index for measuring whether data exhibit phylogenetic dependence or not and whether it has low rates of Type I error. Moreover, λ is robust to incomplete phylogenetic information, which demonstrates that even partial information on phylogeny will improve the accuracy of phylogenetic analyses. To assess whether traits generally show phylogenetic associations, we present a quantitative review of 26 published phylogenetic comparative data sets. The data sets include 103 traits and were chosen from the ecological literature in which debate about the need for phylogenetic correction has been most acute. Eighty‐eight percent of data sets contained at least one character that displayed significant phylogenetic dependence, and 60% of characters overall (pooled across studies) showed significant evidence of phylogenetic association. In 16% of tests, phylogenetic correlation could be neither supported nor rejected. However, most of these equivocal results were found in small phylogenies and probably reflect a lack of power. We suggest that the parameter λ be routinely estimated when analyzing comparative data, since it can also be used simultaneously to adjust the phylogenetic correction in a manner that is optimal for the data set, and we present an example of how this may be done.

What Is the Allee Effect

W. C. Allee brought attention to the possibility of a positive relationship between aspects of fitness and population size over fifty years ago. This phenomenon, frequently termed the Allee effect, has been the focus of increased interest over the past two decades in the light of concerns over conservation and the problems of rarity. Use of the term suffers from the absence of a clear definition however, with the result that Allee effects are frequently thought to involve only a narrow range of phenomena and are often overlooked altogether. We propose a definition for the effect and attempt to resolve the major issues underlying the confusion surrounding this term.

The role of ecological theory in microbial ecology

Microbial ecology is currently undergoing a revolution, with repercussions spreading throughout microbiology, ecology and ecosystem science. The rapid accumulation of molecular data is uncovering vast diversity, abundant uncultivated microbial groups and novel microbial functions. This accumulation of data requires the application of theory to provide organization, structure, mechanistic insight and, ultimately, predictive power that is of practical value, but the application of theory in microbial ecology is currently very limited. Here we argue that the full potential of the ongoing revolution will not be realized if research is not directed and driven by theory, and that the generality of established ecological theory must be tested using microbial systems.

The Origins of C4 Grasslands: Integrating Evolutionary and Ecosystem Science

Grassland Emergence The evolution of the C4 photosynthetic pathway from the ancestral C3 pathway in grasses led to the establishment of grasslands in warm climates during the Late Miocene (8 to 3 million years ago). This was a major event in plant evolutionary history, and their high rates of foliage production sustained high levels of herbivore consumption. The past decade has seen significant advances in understanding C4 grassland ecosystem ecology, and now a wealth of data on the geological history of these ecosystems has accumulated and the phylogeny of grasses is much better known. Edwards et al. (p. 587) review this multidisciplinary research area and attempt to synthesize emerging knowledge about the evolution of grass species within the context of plant and ecosystem ecology. The evolution of grasses using C4 photosynthesis and their sudden rise to ecological dominance 3 to 8 million years ago is among the most dramatic examples of biome assembly in the geological record. A growing body of work suggests that the patterns and drivers of C4 grassland expansion were considerably more complex than originally assumed. Previous research has benefited substantially from dialog between geologists and ecologists, but current research must now integrate fully with phylogenetics. A synthesis of grass evolutionary biology with grassland ecosystem science will further our knowledge of the evolution of traits that promote dominance in grassland systems and will provide a new context in which to evaluate the relative importance of C4 photosynthesis in transforming ecosystems across large regions of Earth.

Identification of 100 fundamental ecological questions

Summary 1. Fundamental ecological research is both intrinsically interesting and provides the basic knowledge required to answer applied questions of importance to the management of the natural world. The 100th anniversary of the British Ecological Society in 2013 is an opportune moment to reflect on the current status of ecology as a science and look forward to high-light priorities for future work.

Pathogens and insect herbivores drive rainforest plant diversity and composition

Tropical forests are important reservoirs of biodiversity, but the processes that maintain this diversity remain poorly understood. The Janzen–Connell hypothesis suggests that specialized natural enemies such as insect herbivores and fungal pathogens maintain high diversity by elevating mortality when plant species occur at high density (negative density dependence; NDD). NDD has been detected widely in tropical forests, but the prediction that NDD caused by insects and pathogens has a community-wide role in maintaining tropical plant diversity remains untested. We show experimentally that changes in plant diversity and species composition are caused by fungal pathogens and insect herbivores. Effective plant species richness increased across the seed-to-seedling transition, corresponding to large changes in species composition. Treating seeds and young seedlings with fungicides significantly reduced the diversity of the seedling assemblage, consistent with the Janzen–Connell hypothesis. Although suppressing insect herbivores using insecticides did not alter species diversity, it greatly increased seedling recruitment and caused a marked shift in seedling species composition. Overall, seedling recruitment was significantly reduced at high conspecific seed densities and this NDD was greatest for the species that were most abundant as seeds. Suppressing fungi reduced the negative effects of density on recruitment, confirming that the diversity-enhancing effect of fungi is mediated by NDD. Our study provides an overall test of the Janzen–Connell hypothesis and demonstrates the crucial role that insects and pathogens have both in structuring tropical plant communities and in maintaining their remarkable diversity.

The seven deadly sins of comparative analysis

Phylogenetic comparative methods are extremely commonly used in evolutionary biology. In this paper, I highlight some of the problems that are frequently encountered in comparative analyses and review how they can be fixed. In broad terms, the problems boil down to a lack of appreciation of the underlying assumptions of comparative methods, as well as problems with implementing methods in a manner akin to more familiar statistical approaches. I highlight that the advent of more flexible computing environments should improve matters and allow researchers greater scope to explore methods and data.

Dealing with collinearity in behavioural and ecological data: model averaging and the problems of measurement error

There has been a great deal of recent discussion of the practice of regression analysis (or more generally, linear modelling) in behaviour and ecology. In this paper, I wish to highlight two factors that have been under-considered, collinearity and measurement error in predictors, as well as to consider what happens when both exist at the same time. I examine what the consequences are for conventional regression analysis (ordinary least squares, OLS) as well as model averaging methods, typified by information theoretic approaches based around Akaike’s information criterion. Collinearity causes variance inflation of estimated slopes in OLS analysis, as is well known. In the presence of collinearity, model averaging reduces this variance for predictors with weak effects, but also can lead to parameter bias. When collinearity is strong or when all predictors have strong effects, model averaging relies heavily on the full model including all predictors and hence the results from this and OLS are essentially the same. I highlight that it is not safe to simply eliminate collinear variables without due consideration of their likely independent effects as this can lead to biases. Measurement error is also considered and I show that when collinearity exists, this can lead to extreme biases when predictors are collinear, have strong effects but differ in their degree of measurement error. I highlight techniques for dealing with and diagnosing these problems. These results reinforce that automated model selection techniques should not be relied on in the analysis of complex multivariable datasets.

Declines in the numbers of amateur and professional taxonomists: implications for conservation

To ensure the effective conservation of biodiversity the distribution of species needs to be accurately characterized and areas of high species richness located. For many taxa this can be achieved only by experienced taxonomists. Taxonomic research has a large input from non-professional or amateur researchers, in addition to professionals working at museums or universities. The decline of taxonomy and the number of taxonomists within the professional community has been widely publicized, but the trends in the activities of amateur taxonomists are unclear. Because amateurs contribute many valuable records of species occurrence this may have a disproportionate impact upon the information available for conservation planning and represents an underappreciated threat to conservation planning. We use taxonomic research by UK entomologists in order to evaluate the changing role of both amateur and professional taxonomists. We reviewed contributions by British-based authors to Entomologists Monthly Magazine over the past century. Our results show that both amateur and professional taxonomy have undergone a long and persistent decline since the 1950s, in terms of both the number of contributors and the number of papers contributed. It is argued that the conservation community needs to help try and reverse the decline of taxonomy.

Space versus phylogeny: disentangling phylogenetic and spatial signals in comparative data.

Variation in traits across species or populations is the outcome of both environmental and historical factors. Trait variation is therefore a function of both the phylogenetic and spatial context of species. Here we introduce a method that, within a single framework, estimates the relative roles of spatial and phylogenetic variations in comparative data. The approach requires traits measured across phylogenetic units, e.g. species, the spatial occurrences of those units and a phylogeny connecting them. The method modifies the expected variance of phylogenetically independent contrasts to include both spatial and phylogenetic effects. We illustrate this approach by analysing cross-species variation in body mass, geographical range size and species-typical environmental temperature in three orders of mammals (carnivores, artiodactyls and primates). These species attributes contain highly disparate levels of phylogenetic and spatial signals, with the strongest phylogenetic autocorrelation in body size and spatial dependence in environmental temperatures and geographical range size showing mixed effects. The proposed method successfully captures these differences and in its simplest form estimates a single parameter that quantifies the relative effects of space and phylogeny. We discuss how the method may be extended to explore a range of models of evolution and spatial dependence.