Statistical ecology comes of age

The desire to predict the consequences of global environmental change has been the driver towards more realistic models embracing the variability and uncertainties inherent in ecology. Statistical ecology has gelled over the past decade as a discipline that moves away from describing patterns towards modelling the ecological processes that generate these patterns. Following the fourth International Statistical Ecology Conference (1–4 July 2014) in Montpellier, France, we analyse current trends in statistical ecology. Important advances in the analysis of individual movement, and in the modelling of population dynamics and species distributions, are made possible by the increasing use of hierarchical and hidden process models. Exciting research perspectives include the development of methods to interpret citizen science data and of efficient, flexible computational algorithms for model fitting. Statistical ecology has come of age: it now provides a general and mathematically rigorous framework linking ecological theory and empirical data.     

Moving beyond abundance distributions: neutral theory and spatial patterns in a tropical forest

Assessing the relative importance of different processes that determine the spatial distribution of species and the dynamics in highly diverse plant communities remains a challenging question in ecology. Previous modelling approaches often focused on single aggregated forest diversity patterns that convey limited information on the underlying dynamic processes. Here, we use recent advances in inference for stochastic simulation models to evaluate the ability of a spatially explicit and spatially continuous neutral model to quantitatively predict six spatial and non-spatial patterns observed at the 50 ha tropical forest plot on Barro Colorado Island, Panama. The patterns capture different aspects of forest dynamics and biodiversity structure, such as annual mortality rate, species richness, species abundance distribution, beta-diversity and the species–area relationship (SAR). The model correctly predicted each pattern independently and up to five patterns simultaneously. However, the model was unable to match the SAR and beta-diversity simultaneously. Our study moves previous theory towards a dynamic spatial theory of biodiversity and demonstrates the value of spatial data to identify ecological processes. This opens up new avenues to evaluate the consequences of additional process for community assembly and dynamics.     

sars: an R package for fitting,evaluating and comparing species–area relationship models

The species–area relationship (SAR) constitutes one of the most general ecological patterns globally. A number of different SAR models have been proposed. Recent work has shown that no single model universally provides the best fit to empirical SAR datasets: multiple models may be of practical and theoretical interest. However, there are no software packages available that a) allow users to fit the full range of published SAR models, or b) provide functions to undertake a range of additional SAR‐related analyses. To address these needs, we have developed the R package ‘sars’ that provides a wide variety of SAR‐related functionality. The package provides functions to: a) fit 20 SAR models using non‐linear and linear regression, b) calculate multi‐model averaged curves using various information criteria, and c) generate confidence intervals using bootstrapping. Plotting functions allow users to depict and scrutinize the fits of individual models and multi‐model averaged curves. The package also provides additional SAR functionality, including functions to fit, plot and evaluate the random placement model using a species–sites abundance matrix, and to fit the general dynamic model of oceanic island biogeography. The ‘sars’ R package will aid future SAR research by providing a comprehensive set of simple to use tools that enable in‐depth exploration of SARs and SAR‐related patterns. The package has been designed to allow other researchers to add new functions and models in the future and thus the package represents a resource for future SAR work that can be built on and expanded by workers in the field.     

Freshwater bacterioplankton richness in oligotrophic lakes depends on nutrient availability rather than on species–area relationships

A central goal in ecology is to grasp the mechanisms that underlie and maintain biodiversity and patterns in its spatial distribution can provide clues about those mechanisms. Here, we investigated what might determine the bacterioplankton richness (BR) in lakes by means of 454 pyrosequencing of the 16S rRNA gene. We further provide a BR estimate based upon a sampling depth and accuracy, which, to our knowledge, are unsurpassed for freshwater bacterioplankton communities. Our examination of 22 669 sequences per lake showed that freshwater BR in fourteen nutrient-poor lakes was positively influenced by nutrient availability. Our study is, thus, consistent with the finding that the supply of available nutrients is a major driver of species richness; a pattern that may well be universally valid to the world of both micro- and macro-organisms. We, furthermore, observed that BR increased with elevated landscape position, most likely as a consequence of differences in nutrient availability. Finally, BR decreased with increasing lake and catchment area that is negative species–area relationships (SARs) were recorded; a finding that re-opens the debate about whether positive SARs can indeed be found in the microbial world and whether positive SARs can in fact be pronounced as one of the few ‘laws'' in ecology.     

Biogeography and ecology: towards the integration of two disciplines

Although ecology and biogeography had common origins in the natural history of the nineteenth century, they diverged substantially during the early twentieth century as ecology became increasingly hypothesis-driven and experimental. This mechanistic focus narrowed ecology''s purview to local scales of time and space, and mostly excluded large-scale phenomena and historical explanations. In parallel, biogeography became more analytical with the acceptance of plate tectonics and the development of phylogenetic systematics, and began to pay more attention to ecological factors that influence large-scale distributions. This trend towards unification exposed problems with terms such as ‘community’ and ‘niche,’ in part because ecologists began to view ecological communities as open systems within the contexts of history and geography. The papers in this issue represent biogeographic and ecological perspectives and address the general themes of (i) the niche, (ii) comparative ecology and macroecology, (iii) community assembly, and (iv) diversity. The integration of ecology and biogeography clearly is a natural undertaking that is based on evolutionary biology, has developed its own momentum, and which promises novel, synthetic approaches to investigating ecological systems and their variation over the surface of the Earth. We offer suggestions on future research directions at the intersection of biogeography and ecology.     

生态学研究的新领域--分子生态学

分子生态是生态学的一个新的研究领域。它采用分子生物学的技术与方法来研究生态活动规律的分子机理。本文简要介绍了分子生态学的定义、研究内容、研究方法和研究热点等,以期描述出分子生态学的概貌。     

A meta-analysis of changes in bacterial and archaeal communities with time

Ecologists have long studied the temporal dynamics of plant and animal communities with much less attention paid to the temporal dynamics exhibited by microbial communities. As a result, we do not know if overarching temporal trends exist for microbial communities or if changes in microbial communities are generally predictable with time. Using microbial time series assessed via high-throughput sequencing, we conducted a meta-analysis of temporal dynamics in microbial communities, including 76 sites representing air, aquatic, soil, brewery wastewater treatment, human- and plant-associated microbial biomes. We found that temporal variability in both within- and between-community diversity was consistent among microbial communities from similar environments. Community structure changed systematically with time in less than half of the cases, and the highest rates of change were observed within ranges of 1 day to 1 month for all communities examined. Microbial communities exhibited species–time relationships (STRs), which describe the accumulation of new taxa to a community, similar to those observed previously for plant and animal communities, suggesting that STRs are remarkably consistent across a broad range of taxa. These results highlight that a continued integration of microbial ecology into the broader field of ecology will provide new insight into the temporal patterns of microbial and ‘macro''-bial communities alike.     

Subjecting the theory of the small‐island effect to Ockham’s razor

Species–area curves from islands and other isolates often differ in shape from sample‐area curves generated from mainlands or sections of isolates (or islands), especially at finer scales. We examine two explanations for this difference: (1) the small‐island effect (SIE), which assumes the species–area curve is composed of two distinctly different curve patterns; and (2) a sigmoid or depressed isolate species–area curve with no break‐points (in arithmetic space). We argue that the application of Ockham’s razor – the principle that the simplest, most economical explanation for a hypothesis should be accepted over less parsimonious alternatives – leads to the conclusion that the latter explanation is preferable. We hold that there is no reason to assume the ecological factors or patterns that affect the shapes of isolate (or island) curves cause two distinctly different patterns. This assumption is not required for the alternative, namely that these factors cause a single (though depressed) isolate species–area curve with no break‐points. We conclude that the theory of the small‐island effect, despite its present standing as an accepted general pattern in nature, should be abandoned.     

The effect of inappropriate calibration: three case studies in molecular ecology

Time-scales estimated from sequence data play an important role in molecular ecology. They can be used to draw correlations between evolutionary and palaeoclimatic events, to measure the tempo of speciation, and to study the demographic history of an endangered species. In all of these studies, it is paramount to have accurate estimates of time-scales and substitution rates. Molecular ecological studies typically focus on intraspecific data that have evolved on genealogical scales, but often these studies inappropriately employ deep fossil calibrations or canonical substitution rates (e.g., 1% per million years for birds and mammals) for calibrating estimates of divergence times. These approaches can yield misleading estimates of molecular time-scales, with significant impacts on subsequent evolutionary and ecological inferences. We illustrate this calibration problem using three case studies: avian speciation in the late Pleistocene, the demographic history of bowhead whales, and the Pleistocene biogeography of brown bears. For each data set, we compare the date estimates that are obtained using internal and external calibration points. In all three cases, the conclusions are significantly altered by the application of revised, internally-calibrated substitution rates. Collectively, the results emphasise the importance of judicious selection of calibrations for analyses of recent evolutionary events.     

A new approach to interspecific synchrony in population ecology using tail association

Standard methods for studying the association between two ecologically important variables provide only a small slice of the information content of the association, but statistical approaches are available that provide comprehensive information. In particular, available approaches can reveal tail associations, that is, accentuated or reduced associations between the more extreme values of variables. We here study the nature and causes of tail associations between phenological or population‐density variables of co‐located species, and their ecological importance. We employ a simple method of measuring tail associations which we call the partial Spearman correlation. Using multidecadal, multi‐species spatiotemporal datasets on aphid first flights and marine phytoplankton population densities, we assess the potential for tail association to illuminate two major topics of study in community ecology: the stability or instability of aggregate community measures such as total community biomass and its relationship with the synchronous or compensatory dynamics of the community''s constituent species; and the potential for fluctuations and trends in species phenology to result in trophic mismatches. We find that positively associated fluctuations in the population densities of co‐located species commonly show asymmetric tail associations; that is, it is common for two species’ densities to be more correlated when large than when small, or vice versa. Ordinary measures of association such as correlation do not take this asymmetry into account. Likewise, positively associated fluctuations in the phenology of co‐located species also commonly show asymmetric tail associations. We provide evidence that tail associations between two or more species’ population‐density or phenology time series can be inherited from mutual tail associations of these quantities with an environmental driver. We argue that our understanding of community dynamics and stability, and of phenologies of interacting species, can be meaningfully improved in future work by taking into account tail associations.     

A primer of community ecology using the R language

Community ecology beginners often struggle to understand theories expressed in complex mathematical formulas and to master computer programming. To remedy this situation, this article provides a practical, R-based introduction to community ecology by illustrating core concepts (vital rates, carrying capacity, and density dependence) and models that can be used to explore the patterns of species abundance and diversity. The structure of this article consists of three modeling exercises, each asking a general question that can be answered by a combination of theory and R programming: (1) what determines the abundance of species, and what makes a population persist and go extinct?; (2) what determines the distribution of species and species diversity?; (3) what determines the relative abundance of species and what allows species to coexist? Through the exercises, I discuss the following five concepts and ideas that provide valuable insights into the questions: (i) the tragedy of the commons, (ii) the theory of island biogeography, (iii) competitive exclusion, (iv) the neutral theory of biodiversity, and (v) frequency dependence. These materials are thus designed to guide the reader in developing an intuition for ecological thinking that will help capture the essence of the global environmental and biodiversity crisis. Although this article does not delineate the scope and depth of the vast field of community ecology, I hope that it will motivate the reader to step up to a more formal introduction to community ecology.     

How Do Snails Meet Fish? Landscape Perspective Needed to Study Parasite Prevalence

We echo the viewpoints presented in the recent editorials and forums in EcoHealth in 2009 and 2010 that understanding the links between water and health using biogeography can provide insights into the patterns of and the processes that give rise to the distribution of disease prevalence. In particular, we underscore the need to integrate disease ecology and biogeography using landscape ecological approaches. We use opisthorchiasis, a major public health problem in Southeast Asia, to illustrate our opinions.     

The evolution and ecology of masquerade

Many organisms appear to mimic inanimate objects such as twigs, leaves, stones, and bird droppings. Such adaptations are considered to have evolved because their bearers are misidentified as either inedible objects by their predators, or as innocuous objects by their prey. In the past, this phenomenon has been classified by some as Batesian mimicry and by others as crypsis, but now is considered to be conceptually different from both, and has been termed ‘masquerade’. Despite the debate over how to classify masquerade, this phenomenon has received little attention from evolutionary biologists. Here, we discuss the limited empirical evidence supporting the idea that masquerade functions to cause misidentification of organisms, provide a testable definition of masquerade, and suggest how masquerade evolved and under what ecological conditions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 1–8.     

景观生态学在海岸带地区的研究进展

海岸带是景观生态学研究的重要领域,并且取得了长足的发展。系统剖析景观生态学在海岸带陆地土地利用变化的景观格局分析与优化、滨海湿地景观格局破碎化及其生态效应、海岛开发和保护的景观规划与设计、近岸海域空间开发的景观格局分析与评价、海洋环境过程与空间格局的相互作用关系等领域研究取得的成绩及其存在的主要问题。在此基础上,结合我国生态文明建设要求,探讨了景观生态学在海洋生态文明建设中的研究尺度匹配、格局-过程耦合理论引导、景观格局指数量化考核方面的学科优势。景观生态学可以在围填海平面设计与规划研究、海岸带生态修复与景观建设、海洋保护区选划与管理、海岛开发与保护、海陆统筹下的海岸带综合管理等方面开拓应用发展领域,助力海洋生态文明建设,承担建设海洋强国、美丽海洋的新使命。同时也为景观生态学在近岸海洋资源环境中的应用、近岸海洋景观生态学的发展梳理框架与思路。     

The use of Hardy–Weinberg Equilibrium in clonal plant systems

Traditionally population genetics precludes the use of the same genetic individual more than once in Hardy–Weinberg (HW) based calculations due to the model''s explicit assumptions. However, when applied to clonal plant populations this can be difficult to do, and in some circumstances, it may be ecologically informative to use the ramet as the data unit. In fact, ecologists have varied the definition of the individual from a strict adherence to a single data point per genotype to a more inclusive approach of one data point per ramet. With the advent of molecular tools, the list of facultatively clonal plants and the recognition of their ecological relevance grows. There is an important risk of misinterpretation when HW calculations are applied to a clonal plant not recognized as clonal, as well as when the definition of the individual for those calculations is not clearly stated in a known clonal species. Focusing on heterozygosity values, we investigate cases that demonstrate the extreme range of potential modeling outcomes and describe the different contexts where a particular definition could better meet ecological modeling goals. We emphasize that the HW model can be ecologically relevant when applied to clonal plants, but caution is necessary in how it is used, reported, and interpreted. We propose that in known clonal plants, both genotype (GHet) and ramet (RHet) based calculations are reported to define the full range of potential values and better facilitate cross‐study comparisons.     

Biogeography and ecology: two views of one world

Both biogeography and ecology seek to understand the processes that determine patterns in nature, but do so at different spatial and temporal scales. The two disciplines were not always so different, and are recently converging again at regional spatial scales and broad temporal scales. In order to avoid confusion and to hasten progress at the converging margins of each discipline, the following papers were presented at a symposium in the International Biogeography Society''s 2011 meeting, and are now published in this issue of the Philosophical Transactions of the Royal Society B. In a novel approach, groups of authors were paired to represent biogeographic and ecological perspectives on each of four topics: niche, comparative ecology and macroecology, community assembly, and diversity. Collectively, this compilation identifies points of agreement and disagreement between the two views on these central topics, and points to future research directions that may build on agreements and reconcile differences. We conclude this compilation with an overview on the integration of biogeography and ecology.     

Historical biogeography, ecology and species richness

Ecology and historical (phylogeny-based) biogeography have much to offer one another, but exchanges between these fields have been limited. Historical biogeography has become narrowly focused on using phylogenies to discover the history of geological connections among regions. Conversely, ecologists often ignore historical biogeography, even when its input can be crucial. Both historical biogeographers and ecologists have more-or-less abandoned attempts to understand the processes that determine the large-scale distribution of clades. Here, we describe the chasm that has developed between ecology and historical biogeography, some of the important questions that have fallen into it and how it might be bridged. To illustrate the benefits of an integrated approach, we expand on a model that can help explain the latitudinal gradient of species richness.     

The relationship between biogeography and ecology: envelopes,models, predictions

This paper reviews ideas on the relationship between the ecology of clades and their distribution. Ecological biogeography represents a tradition that dates back to ancient times. It assumes that the distribution of organisms is explained by factors of present environment, especially climate. In contrast, modern systematics, following its origins in the Renaissance, concluded with Darwin that ‘neither the similarity nor the dissimilarity of the inhabitants of various regions can be accounted for by their climatal and other physical conditions’. In many cases, species distribution models – ecological niche models – based on the current environment of a species (its environmental envelope) fail to predict the actual distribution of the species. In particular, they often over‐predict distributions. In addition, a group's niche often varies in space and time. These results provide valuable evidence that Darwin was correct, and many ecologists now recognise that there is a problem with the niche theory of distribution. Current ecological processes explain distribution at smaller scales than do biogeographical and evolutionary processes, but the latter can lead to patterns that are much more local than many ecologists have assumed. Biogeographical phenomena often occur at a much smaller scale than that of the Wallacean regions. In areas that have been subjected to marine inundation or intense tectonism, many centres of endemism are only tens of kilometres across. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 456–468.     

Using ecological networks to answer questions in global biogeography and ecology

Ecological networks have classically been studied at site and landscape scales, yet recent efforts have been made to collate these data into global repositories. This offers an opportunity to integrate and upscale knowledge about ecological interactions from local to global scales to gain enhanced insights from the mechanistic information provided by these data. By drawing on existing research investigating patterns in ecological interactions at continental to global scales, we show how data on ecological networks, collected at appropriate scales, can be used to generate an improved understanding of many aspects of ecology and biogeography—for example, species distribution modelling, restoration ecology and conservation. We argue that by understanding the patterns in the structure and function of ecological networks across scales, it is possible to enhance our understanding of the natural world.