Predicting species abundance distributions by simultaneously using number and biomass as units of measurement

The universal observation that some species in an ecological community are common, but many more are rare, is neatly encapsulated in a species abundance distribution (SAD)1. However, the shape of the distribution can depend on the currency used to measure abundance 2. Here we show how the SADs for numerical abundance and biomass are related and how this relationship can be used to predict the form of the SAD. When plotted in log numerical abundance, log biomass space, species points lie within an approximately triangular area the limits of which are set by body size range, and the upper limit of abundance in both metrics. Under the simplifying, but reasonable, assumption that the observed scatter of species within this region is random, the shape of the SAD is immediately derived from simple geometrical considerations. For the SAD of numerical abundance this is a power curve. The biomass SAD can be either a power curve or, more frequently, a unimodal curve, which can approximate a log normal. This log triangular random placement model serves as a null hypothesis against which actual communities can be compared. Data from two intensively surveyed local communities indicate that it can give a good approximation, with species scattered within a triangle. Further, we can predict the consequences, for the SAD, of size-selective sampling protocols. We argue that mechanistic models of SADs must be able to account for the relative abundance of species in alternative currencies. Moreover, this approach will shed light on niche packing and may have application in environmental monitoring

Recent increases in assemblage rarity are linked to increasing local immigration

F.A.M.J.'s PhD was financed by the School of Biology, University of St Andrews. M.D. and A.E.M. acknowledge funding by the Leverhulme Trust. A.E.M. acknowledges funding from the European Research Council (ERC AdG BioTIME 250189 and ERC PoC BioCHANGE 727440).As pressures on biodiversity increase, a better understanding of how assemblages are responding is needed. Because rare species, defined here as those that have locally low abundances, make up a high proportion of assemblage species lists, understanding how the number of rare species within assemblages is changing will help elucidate patterns of recent biodiversity change. Here, we show that the number of rare species within assemblages is increasing, on average, across systems. This increase could arise in two ways: species already present in the assemblage decreasing in abundance but with no increase in extinctions, or additional species entering the assemblage in low numbers associated with an increase in immigration. The positive relationship between change in rarity and change in species richness provides evidence for the second explanation, i.e. higher net immigration than extinction among the rare species. These measurable changes in the structure of assemblages in the recent past underline the need to use multiple biodiversity metrics to understand biodiversity change.Publisher PDFPeer reviewe

Complex community responses underpin biodiversity change following invasion

How do invasive species change native biodiversity? One reason why this long-standing question remains challenging to answer could be because the main focus of the invasion literature has been on shifts in species richness (a measure of α-diversity). As the underlying components of community structure—intraspecific aggregation, interspecific density and the species abundance distribution (SAD)—are potentially impacted in different ways during invasion, trends in species richness provide only limited insight into the mechanisms leading to biodiversity change. In addition, these impacts can be manifested in distinct ways at different spatial scales. Here we take advantage of the new Measurement of Biodiversity (MoB) framework to reanalyse data collected in an invasion front in the Brazilian Cerrado biodiversity hotspot. We show that, by using the MoB multi-scale approach, we are able to link reductions in species richness in invaded sites to restructuring in the SAD. This restructuring takes the form of lower evenness in sites invaded by pines relative to sites without pines. Shifts in aggregation also occur. There is a clear signature of spatial scale in biodiversity change linked to the presence of an invasive species. These results demonstrate how the MoB approach can play an important role in helping invasion ecologists, field biologists and conservation managers move towards a more mechanistic approach to detecting and interpreting changes in ecological systems following invasion

Goodness-of-fit measures of evenness: a new tool for exploring changes in community structure

Growing concern about the fate of biodiversity, highlighted by the Convention on Biological Diversity's 2010 and 2020 targets for stemming biodiversity loss, has intensified interest in methods of assessing change in ecological communities through time. Biodiversity is a multivariate concept, which cannot be well‐represented by a single measure. However, diversity profiles summarize the multivariate nature of multi‐species datasets, and allow a more nuanced interpretation of biodiversity trends than unitary metrics. Here we introduce a new approach to diversity profiling. Our method is based on the knowledge that an ecological community is never completely even and uses this departure from perfect evenness as a novel and insightful way of measuring diversity. We plot our measure of departure as a function of a free parameter, to generate “evenness profiles”. These profiles allow us to separate changes due to dominant species from those due to rare species, and relate these patterns to shifts in overall diversity. This separation of the influence of dominance and rarity on overall diversity enables the user to uncover changes in diversity that would be masked in other methods. We discuss profiling techniques based on this parametric family, and explore its connections with existing diversity indices. Next, we evaluate our approach in terms of predicted community structure (following Tokeshi's niche models) and present an example assessing temporal trends in diversity of British farmland birds. We conclude that this method is an informative and tractable parametric approach for quantifying evenness. It provides novel insights into community structure, revealing the contributions of both rare and common species to biodiversity trends

Temporal turnover and the maintenance of diversity in ecological assemblages

Temporal variation in species abundances occurs in all ecological communities. Here, we explore the role that this temporal turnover plays in maintaining assemblage diversity. We investigate a three-decade time series of estuarine fishes and show that the abundances of the individual species fluctuate asynchronously around their mean levels. We then use a time-series modelling approach to examine the consequences of different patterns of turnover, by asking how the correlation between the abundance of a species in a given year and its abundance in the previous year influences the structure of the overall assemblage. Classical diversity measures that ignore species identities reveal that the observed assemblage structure will persist under all but the most extreme conditions. However, metrics that track species identities indicate a narrower set of turnover scenarios under which the predicted assemblage resembles the natural one. Our study suggests that species diversity metrics are insensitive to change and that measures that track species ranks may provide better early warning that an assemblage is being perturbed. It also highlights the need to incorporate temporal turnover in investigations of assemblage structure and function

The impact of resource dependence of the mechanisms of life on the spatial population dynamics of an in silico microbial community

Biodiversity has a critical impact on ecosystem functionality and stability, and thus the current biodiversity crisis has motivated many studies of the mechanisms that sustain biodiversity, a notable example being non-transitive or cyclic competition. We therefore extend existing microscopic models of communities with cyclic competition by incorporating resource dependence in demographic processes, characteristics of natural systems often oversimplified or overlooked by modellers. The spatially explicit nature of our individual-based model of three interacting species results in the formation of stable spatial structures, which have significant effects on community functioning, in agreement with experimental observations of pattern formation in microbial communities. Published by AIP Publishing

Inferring macro-ecological patterns from local presence/absence data

Biodiversity provides support for life, vital provisions, regulating services and has positive cultural impacts. It is therefore important to have accurate methods to measure biodiversity, in order to safeguard it when we discover it to be threatened. For practical reasons, biodiversity is usually measured at fine scales whereas diversity issues (e.g. conservation) interest regional or global scales. Moreover, biodiversity may change across spatial scales. It is therefore a key challenge to be able to translate local information on biodiversity into global patterns. Many databases give no information about the abundances of a species within an area, but only its occurrence in each of the surveyed plots. In this paper, we introduce an analytical framework (implemented in a ready‐to‐use R code) to infer species richness and abundances at large spatial scales in biodiversity‐rich ecosystems when species presence/absence information is available on various scattered samples (i.e. upscaling). This framework is based on the scale‐invariance property of the negative binomial. Our approach allows to infer and link within a unique framework important and well‐known biodiversity patterns of ecological theory, such as the species accumulation curve (SAC) and the relative species abundance (RSA) as well as a new emergent pattern, which is the relative species occupancy (RSO). Our estimates are robust and accurate, as confirmed by tests performed on both in silico‐generated and real forests. We demonstrate the accuracy of our predictions using data from two well‐studied forest stands. Moreover, we compared our results with other popular methods proposed in the literature to infer species richness from presence to absence data and we showed that our framework gives better estimates. It has thus important applications to biodiversity research and conservation practice

More than the sum of the parts : annual partitioning within spatial guilds underpins community regulation

Funding: ERC (BioTIME 250189 and BioCHANGE 727440) and the Royal Society.To withstand the pressures of a rapidly changing world, resilient ecosystems should exhibit compensatory dynamics, including uncorrelated temporal shifts in population sizes. The observation that diversity is maintained through time in many systems is evidence that communities are indeed regulated and stabilized, yet empirical observations suggest that positive covariance in species abundances is widespread. This paradox could be resolved if communities are composed of a number of ecologically relevant sub-units in which the members compete for resources, but whose abundances fluctuate independently. Such modular organization could explain community regulation, even when the community as a whole appears synchronized.To test this hypothesis, we quantified temporal synchronicity in annual population abundances within spatial guilds in an estuarine fish assemblage that has been monitored for 36 years. We detected independent fluctuations in annual abundances within guilds. By contrast, the assemblage as a whole exhibited temporal synchronicity—an outcome linked to the dynamics of guild dominants, which were synchronized with each other. These findings underline the importance of modularity in explaining community regulation and highlight the need to protect assemblage composition and structure as well as species richness.PostprintPeer reviewe

Dominance structure of assemblages is regulated over a period of rapid environmental change

F.A.M.J. is financed by the School of Biology, University of St Andrews. A.E.M. acknowledges funding from the European Research Council (ERCAdG BioTIME 250189 and ERCPoC BioCHANGE 727440).Ecological assemblages are inherently uneven, with numerically dominant species contributing disproportionately to ecosystem services. Marked biodiversity change due to growing pressures on the world's ecosystems is now well documented. However, the hypothesis that dominant species are becoming relatively more abundant has not been tested. We examined the prediction that the dominance structure of contemporary communities is shifting, using a meta-analysis of 110 assemblage timeseries. Changes in relative and absolute dominance were evaluated with mixed and cyclic-shift permutation models. Our analysis uncovered no evidence of a systematic change in either form of dominance, but established that relative dominance is preserved even when assemblage size (total N) changes. This suggests that dominance structure is regulated alongside richness and assemblage size, and highlights the importance of investigating multiple components of assemblage diversity when evaluating ecosystem responses to environmental drivers.PostprintPeer reviewe

Measuring temporal change in alpha diversity : a framework integrating taxonomic, phylogenetic and functional diversity and the iNEXT.3D standardization

Funding: This work is jointly supported by the Natural Environment Research Council, UK (NE/T004487/1 for AM and MD) and the Taiwan Ministry of Science and Technology under Contracts NERC-MOST 108-2923-M-007-003 (for AC and CC). AM and MD also acknowledge support from the Leverhulme Trust (RPG-2019-401).1. Biodiversity is a multifaceted concept covering different levels of organisation from genes to ecosystems. Biodiversity has at least three dimensions: (i) Taxonomic diversity (TD): a measure that is sensitive to the number and abundances of species. (ii) Phylogenetic diversity (PD): a measure that incorporates not only species abundances but also species evolutionary histories. (iii) Functional diversity (FD): a measure that considers not only species abundances but also species? traits. 2. We integrate the three dimensions of diversity under a unified framework of Hill numbers and their generalizations. Our TD quantifies the effective number of equally-abundant species, PD quantifies the effective total branch length, mean-PD (PD divided by tree depth) quantifies the effective number of equally-divergent lineages, and FD quantifies the effective number of equally-distinct virtual functional groups (or functional ?species?). Thus, TD, mean-PD and FD are all in the same units of species/lineage equivalents and can be meaningfully compared. 3. Like species richness, empirical TD, PD and FD based on sampling data, depend on sampling effort and sample completeness. For TD (Hill numbers), the iNEXT (interpolation and extrapolation) standardization was developed for standardizing sample size or sample completeness (as measured by sample coverage, the fraction of individuals that belong to the observed species) to make objective comparisons across studies. This paper extends the iNEXT method to the iNEXT.3D standardization to encompass all three dimensions of diversity via sample-size- and sample-coverage-based rarefaction and extrapolation under the unified framework. The asymptotic diversity estimates (i.e., sample size tends to infinity and sample coverage tends to unity) are also derived. In addition to individual-based abundance data, the proposed iNEXT.3D standardization is adapted to deal with incidence-based occurrence data. 4. We apply the integrative framework and the proposed iNEXT.3D standardization to measure temporal alpha-diversity changes for estuarine fish assemblage data spanning four decades. The influence of environmental drivers on diversity change are also assessed. Our analysis informs a mechanistic interpretation of biodiversity change in the three dimensions of diversity. The accompanying freeware, iNEXT.3D, developed during this project, facilitates all computation and graphics.PostprintPeer reviewe