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Niche theory predicts that ecologically similar species can coexist through multidimensional niche partitioning. However, owing to the challenges of accounting for both abiotic and biotic processes in ecological niche modelling, the underlying mechanisms that facilitate coexistence of competing species are poorly understood. In this study, we evaluated potential mechanisms underlying the coexistence of ecologically similar bird species in a biodiversity-rich transboundary montane forest in east-central Africa by computing niche overlap indices along an environmental elevation gradient, diet, forest strata, activity patterns and within-habitat segregation across horizontal space. We found strong support for abiotic environmental habitat niche partitioning, with 55% of species pairs having separate elevation niches. For the remaining species pairs that exhibited similar elevation niches, we found that within-habitat segregation across horizontal space and to a lesser extent vertical forest strata provided the most likely mechanisms of species coexistence. Coexistence of ecologically similar species within a highly diverse montane forest was determined primarily by abiotic factors (e.g. environmental elevation gradient) that characterize the Grinnellian niche and secondarily by biotic factors (e.g. vertical and horizontal segregation within habitats) that describe the Eltonian niche. Thus, partitioning across multiple levels of spatial organization is a key mechanism of coexistence in diverse communities.more » « less
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Abstract Numerous modelling techniques exist to estimate abundance of plant and animal populations. The most accurate methods account for multiple complexities found in ecological data, such as observational biases, spatial autocorrelation, and species correlations. There is, however, a lack of user‐friendly and computationally efficient software to implement the various models, particularly for large data sets.
We developed the
spAbundance R package for fitting spatially explicit Bayesian single‐species and multi‐species hierarchical distance sampling models, N‐mixture models, and generalized linear mixed models. The models within the package can account for spatial autocorrelation using Nearest Neighbour Gaussian Processes and accommodate species correlations in multi‐species models using a latent factor approach, which enables model fitting for data sets with large numbers of sites and/or species.We provide three vignettes and three case studies that highlight
spAbundance functionality. We used spatially explicit multi‐species distance sampling models to estimate density of 16 bird species in Florida, USA, an N‐mixture model to estimate black‐throated blue warbler (Setophaga caerulescens ) abundance in New Hampshire, USA, and a spatial linear mixed model to estimate forest above‐ground biomass across the continental USA.spAbundance provides a user‐friendly, formula‐based interface to fit a variety of univariate and multivariate spatially explicit abundance models. The package serves as a useful tool for ecologists and conservation practitioners to generate improved inference and predictions on the spatial drivers of abundance in populations and communities. -
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Abstract Determining the spatial distributions of species and communities is a key task in ecology and conservation efforts. Joint species distribution models are a fundamental tool in community ecology that use multi‐species detection–nondetection data to estimate species distributions and biodiversity metrics. The analysis of such data is complicated by residual correlations between species, imperfect detection, and spatial autocorrelation. While many methods exist to accommodate each of these complexities, there are few examples in the literature that address and explore all three complexities simultaneously. Here we developed a spatial factor multi‐species occupancy model to explicitly account for species correlations, imperfect detection, and spatial autocorrelation. The proposed model uses a spatial factor dimension reduction approach and Nearest Neighbor Gaussian Processes to ensure computational efficiency for data sets with both a large number of species (e.g., >100) and spatial locations (e.g., 100,000). We compared the proposed model performance to five alternative models, each addressing a subset of the three complexities. We implemented the proposed and alternative models in the
spOccupancy software, designed to facilitate application via an accessible, well documented, and open‐source R package. Using simulations, we found that ignoring the three complexities when present leads to inferior model predictive performance, and the impacts of failing to account for one or more complexities will depend on the objectives of a given study. Using a case study on 98 bird species across the continental US, the spatial factor multi‐species occupancy model had the highest predictive performance among the alternative models. Our proposed framework, together with its implementation inspOccupancy , serves as a user‐friendly tool to understand spatial variation in species distributions and biodiversity while addressing common complexities in multi‐species detection–nondetection data. -
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Abstract Aim Species distribution models (SDMs) are increasingly applied across macroscales using detection‐nondetection data. These models typically assume that a single set of regression coefficients can adequately describe species–environment relationships and/or population trends. However, such relationships often show nonlinear and/or spatially varying patterns that arise from complex interactions with abiotic and biotic processes that operate at different scales. Spatially varying coefficient (SVC) models can readily account for variability in the effects of environmental covariates. Yet, their use in ecology is relatively scarce due to gaps in understanding the inferential benefits that SVC models can provide compared to simpler frameworks.
Innovation Here we demonstrate the inferential benefits of SVC SDMs, with a particular focus on how this approach can be used to generate and test ecological hypotheses regarding the drivers of spatial variability in population trends and species–environment relationships. We illustrate the inferential benefits of SVC SDMs with simulations and two case studies: one that assesses spatially varying trends of 51 forest bird species in the eastern United States over two decades and a second that evaluates spatial variability in the effects of five decades of land cover change on grasshopper sparrow (
Ammodramus savannarum ) occurrence across the continental United States.Main conclusions We found strong support for SVC SDMs compared to simpler alternatives in both empirical case studies. Factors operating at fine spatial scales, accounted for by the SVCs, were the primary divers of spatial variability in forest bird occurrence trends. Additionally, SVCs revealed complex species–habitat relationships with grassland and cropland area for grasshopper sparrow, providing nuanced insights into how future land use change may shape its distribution. These applications display the utility of SVC SDMs to help reveal the environmental factors that drive species distributions across both local and broad scales. We conclude by discussing the potential applications of SVC SDMs in ecology and conservation.
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Abstract Data deficiencies among rare or cryptic species preclude assessment of community‐level processes using many existing approaches, limiting our understanding of the trends and stressors for large numbers of species. Yet evaluating the dynamics of whole communities, not just common or charismatic species, is critical to understanding and the responses of biodiversity to ongoing environmental pressures.
A recent surge in both public science and government‐funded data collection efforts has led to a wealth of biodiversity data. However, these data collection programmes use a wide range of sampling protocols (from unstructured, opportunistic observations of wildlife to well‐structured, design‐based programmes) and record information at a variety of spatiotemporal scales. As a result, available biodiversity data vary substantially in quantity and information content, which must be carefully reconciled for meaningful ecological analysis.
Hierarchical modelling, including single‐species integrated models and hierarchical community models, has improved our ability to assess and predict biodiversity trends and processes. Here, we highlight the emerging ‘integrated community modelling’ framework that combines both data integration and community modelling to improve inferences on species‐ and community‐level dynamics.
We illustrate the framework with a series of worked examples. Our three case studies demonstrate how integrated community models can be used to extend the geographic scope when evaluating species distributions and community‐level richness patterns; discern population and community trends over time; and estimate demographic rates and population growth for communities of sympatric species. We implemented these worked examples using multiple software methods through the R platform via packages with formula‐based interfaces and through development of custom code in JAGS, NIMBLE and Stan.
Integrated community models provide an exciting approach to model biological and observational processes for multiple species using multiple data types and sources simultaneously, thus accounting for uncertainty and sampling error within a unified framework. By leveraging the combined benefits of both data integration and community modelling, integrated community models can produce valuable information about both common and rare species as well as community‐level dynamics, allowing for holistic evaluation of the effects of global change on biodiversity.
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spOccupancy: An R package for single‐species, multi‐species, and integrated spatial occupancy modelsmore » « less
Abstract Occupancy modelling is a common approach to assess species distribution patterns, while explicitly accounting for false absences in detection–nondetection data. Numerous extensions of the basic single‐species occupancy model exist to model multiple species, spatial autocorrelation and to integrate multiple data types. However, development of specialized and computationally efficient software to incorporate such extensions, especially for large datasets, is scarce or absent.
We introduce the
spOccupancy R package designed to fit single‐species and multi‐species spatially explicit occupancy models. We fit all models within a Bayesian framework using Pólya‐Gamma data augmentation, which results in fast and efficient inference.spOccupancy provides functionality for data integration of multiple single‐species detection–nondetection datasets via a joint likelihood framework. The package leverages Nearest Neighbour Gaussian Processes to account for spatial autocorrelation, which enables spatially explicit occupancy modelling for potentially massive datasets (e.g. 1,000s–100,000s of sites).spOccupancy provides user‐friendly functions for data simulation, model fitting, model validation (by posterior predictive checks), model comparison (using information criteria and k‐fold cross‐validation) and out‐of‐sample prediction. We illustrate the package's functionality via a vignette, simulated data analysis and two bird case studies.The
spOccupancy package provides a user‐friendly platform to fit a variety of single and multi‐species occupancy models, making it straightforward to address detection biases and spatial autocorrelation in species distribution models even for large datasets. -
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Abstract 1. The occurrence and distributions of wildlife populations and communities are shifting as a result of global changes. To evaluate whether these shifts are negatively impacting biodiversity processes, it is critical to monitor the status, trends and effects of environmental variables on entire communities. However, modelling the dynamics of multiple species simultaneously can require large amounts of diverse data, and few modelling approaches exist to simultaneously provide species and community‐level inferences.
2. We present an ‘integrated community occupancy model’ (ICOM) that unites principles of data integration and hierarchical community modelling in a single framework to provide inferences on species‐specific and community occurrence dynamics using multiple data sources. The ICOM combines replicated and nonreplicated detection–nondetection data sources using a hierarchical framework that explicitly accounts for different detection and sampling processes across data sources. We use simulations to compare the ICOM to previously developed hierarchical community occupancy models and single species integrated distribution models. We then apply our model to assess the occurrence and biodiversity dynamics of foliage‐gleaning birds in the White Mountain National Forest in the northeastern USA from 2010 to 2018 using three independent data sources.
3. Simulations reveal that integrating multiple data sources in the ICOM increased precision and accuracy of species and community‐level inferences compared to single data source models, although benefits of integration were dependent on the information content of individual data sources (e.g. amount of replication). Compared to single species models, the ICOM yielded more precise species‐level estimates. Within our case study, the ICOM had the highest out‐of‐sample predictive performance compared to single species models and models that used only a subset of the three data sources.
4. The ICOM provides more precise estimates of occurrence dynamics compared to multi‐species models using single data sources or integrated single‐species models. We further found that the ICOM had improved predictive performance across a broad region of interest with an empirical case study of forest birds. The ICOM offers an attractive approach to estimate species and biodiversity dynamics, which is additionally valuable to inform management objectives of both individual species and their broader communities.
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Abstract Monitoring wildlife abundance across space and time is an essential task to study their population dynamics and inform effective management. Acoustic recording units are a promising technology for efficiently monitoring bird populations and communities. While current acoustic data models provide information on the presence/absence of individual species, new approaches are needed to monitor population abundance, ideally across large spatio‐temporal regions.
We present an integrated modelling framework that combines high‐quality but temporally sparse bird point count survey data with acoustic recordings. Our models account for imperfect detection in both data types and false positive errors in the acoustic data. Using simulations, we compare the accuracy and precision of abundance estimates using differing amounts of acoustic vocalizations obtained from a clustering algorithm, point count data, and a subset of manually validated acoustic vocalizations. We also use our modelling framework in a case study to estimate abundance of the Eastern Wood‐Pewee (
Contopus virens ) in Vermont, USA.The simulation study reveals that combining acoustic and point count data via an integrated model improves accuracy and precision of abundance estimates compared with models informed by either acoustic or point count data alone. Improved estimates are obtained across a wide range of scenarios, with the largest gains occurring when detection probability for the point count data is low. Combining acoustic data with only a small number of point count surveys yields estimates of abundance without the need for validating any of the identified vocalizations from the acoustic data. Within our case study, the integrated models provided moderate support for a decline of the Eastern Wood‐Pewee in this region.
Our integrated modelling approach combines dense acoustic data with few point count surveys to deliver reliable estimates of species abundance without the need for manual identification of acoustic vocalizations or a prohibitively expensive large number of repeated point count surveys. Our proposed approach offers an efficient monitoring alternative for large spatio‐temporal regions when point count data are difficult to obtain or when monitoring is focused on rare species with low detection probability.
