A case study on joint species distribution modelling with bird atlas data: revealing limits to species' niches

Growing interest in biodiversity mapping has spurred the development of species distribution atlases, often mainly based on citizen-science projects. Atlas data have been frequently exploited to model species' ecological niches and distributions on a species-by-species basis. However, spatial autocorrelation and phylogenetic relatedness among species complicate the statistical description of species' niches. Also, the effects of species' traits and co-occurrences on species-habitat relationship are commonly disregarded. In this work, we build a hierarchical multi-species model based on a major citizen-science project (the third Spanish breeding bird atlas) that simultaneously accounts for spatial, phylogenetic and trait-based dependencies. We predict the distributions of species niches, species richness and community traits along regional ecological gradients. Climate, habitat associations and species' traits all contribute (in this order) to structuring species' distributions. Species richness increases towards intermediate climatic conditions and with aquatic habitat cover and decreases with increasing forest and woody agricultural land cover. Species were distributed along regional climate gradients in accordance with their global thermal niches. Forest habitats favoured assemblages dominated by generalist, small-sized and cold-dwelling species with limited migratory behaviour. Increasing sampling effort augmented the model performance. Model performance was weaker for rare species and those with decreasing population sizes, likely due to their low niche saturation. Overall, we show that ecological relationships generalize from local to large scales and may be eludicated from atlases based on citizen-science mapping effortsWe wholeheartedly thank JS worked within the REMEDINAL4 network (TE-CM S2018/EMT-4338) during the preparation of the manuscript with no specific funding for this study; he is part of the project Grant (NextDive; PID2021-124187NB-I00) funded by MCIN/AEI/ 10.13039/ 501100011033 and by “ERDF A way of making Europe”. AE was funded by Organismo Autonomo ´ Parques Nacionales of Spain through the project 2745/2021. MJ was supported by the Academy of Finland's ‘Thriving Nature’ profiling action. OO was funded by the Academy of Finland (grant no. 309581), the Jane and Aatos Erkko Foundation, the Research Council of Norway's Centres of Excellence Funding Scheme (223257), and the European Research Council under the Horizon 2020 research and innovation programme (grant agreement No 856506; ERC Synergy project LIFEPLAN). The CSC – IT Center for Science, Finland is acknowledged for computational resource

Contrasting Effects of Chronic Anthropogenic Disturbance on Activity and Species Richness of Insectivorous Bats in Neotropical Dry Forest

For prioritizing conservation actions, it is vital to understand how ecologically diverse species respond to environmental change caused by human activity. This is particularly necessary considering that chronic human disturbance is a threat to biodiversity worldwide. Depending on how species tolerate and adapt to such disturbance, ecological integrity and ecosystem services will be more or less affected. Bats are a species-rich and functionally diverse group, with important roles in ecosystems, and are therefore recognized as a good model group for assessing the impact of environmental change. Their populations have decreased in several regions, especially in the tropics, and are threatened by increasing human disturbance. Using passive acoustic monitoring, we assessed how the species-rich aerial insectivorous bats—essential for insect suppression services—respond to chronic human disturbance in the Caatinga dry forests of Brazil, an area potentially harboring ca. 100 bat species (nearly 50% are insectivorous), but with > 60% its area composed of anthropogenic ecosystems under chronic pressure. Acoustic data for bat activity was collected at research sites with varying amounts of chronic human disturbance (e.g., livestock grazing and firewood gathering). The intensity of the disturbance is indicated by the global multi-metric CAD index (GMDI). Using Animal Sound Identifier (ASI) software, we identified 18 different bat taxon units. Using Hierarchical Modeling of Species Communities (HMSC), we found trends in the association of the disturbance gradient with species richness and bat activity: species richness was higher at sites with higher human disturbance, whereas bat activity decreased with increasing human disturbance. Additionally, we observed taxon-specific responses to human disturbance. We conclude that the effects of chronic anthropogenic disturbance on the insectivorous bat fauna in the Caatinga are not homogeneous and a species-specific approach is necessary when assessing the responses of local bats to human disturbances in tropical dry forests, and in other biomes under human pressure.Peer reviewe

Host--parasite models on graphs

The behavior of two interacting populations, ``hosts''and ``parasites'', is investigated on Cayley trees and scale-free networks. In the former case analytical and numerical arguments elucidate a phase diagram, whose most interesting feature is the absence of a tri-critical point as a function of the two independent spreading parameters. For scale-free graphs, the parasite population can be described effectively by Susceptible-Infected-Susceptible-type dynamics in a host background. This is shown both by considering the appropriate dynamical equations and by numerical simulations on Barab\'asi-Albert networks with the major implication that in the termodynamic limit the critical parasite spreading parameter vanishes.Comment: 10 pages, 6 figures, submitted to PRE; analytics redone, new calculations added, references added, appendix remove

The past and the present in decision-making: the use of conspecific and heterospecific cues in nest site selection

Nest-site selection significantly affects fitness, so adaptations for assessment of the qualities of available sites are expected. The assessment may be based on personal or social information, social information referring to the observed location and performance of both con- and heterospecific individuals. Contrary to large-scale breeding habitat selection, small-scale nest-site selection within habitat patches is insufficiently understood. We analyzed nest-site selection in the migratory collared flycatcher Ficedula albicollis in relation to present and past cues provided by conspecifics and by resident tits within habitat patches by using long-term data. Collared flycatchers preferred nest boxes that were occupied by conspecifics in the previous year. This preference was strongest in breeding pairs where both individuals bred in the same forest patch in the previous year. The results also suggest preference for nest boxes close to boxes where conspecifics had a high breeding success in the previous year, and for nest boxes which are presently surrounded by a high number of breeding great tits Parus major. The results indicate social information use in nest-site selection at a small spatial scale, where collared flycatchers use conspecific cues with a time lag of one year and heterospecific cues instantly.</p

Joint species distribution modelling with the r-package Hmsc

Joint Species Distribution Modelling (JSDM) is becoming an increasingly popular statistical method for analysing data in community ecology. Hierarchical Modelling of Species Communities (HMSC) is a general and flexible framework for fitting JSDMs. HMSC allows the integration of community ecology data with data on environmental covariates, species traits, phylogenetic relationships and the spatio-temporal context of the study, providing predictive insights into community assembly processes from non-manipulative observational data of species communities. The full range of functionality of HMSC has remained restricted to Matlab users only. To make HMSC accessible to the wider community of ecologists, we introduce Hmsc 3.0, a user-friendly r implementation. We illustrate the use of the package by applying Hmsc 3.0 to a range of case studies on real and simulated data. The real data consist of bird counts in a spatio-temporally structured dataset, environmental covariates, species traits and phylogenetic relationships. Vignettes on simulated data involve single-species models, models of small communities, models of large species communities and models for large spatial data. We demonstrate the estimation of species responses to environmental covariates and how these depend on species traits, as well as the estimation of residual species associations. We demonstrate how to construct and fit models with different types of random effects, how to examine MCMC convergence, how to examine the explanatory and predictive powers of the models, how to assess parameter estimates and how to make predictions. We further demonstrate how Hmsc 3.0 can be applied to normally distributed data, count data and presence-absence data. The package, along with the extended vignettes, makes JSDM fitting and post-processing easily accessible to ecologists familiar with r.Peer reviewe

Design matters : an evaluation of the impact of small man-made forest clearings on tropical bats using a before-after-control-impact design

In recent years, large clearings (>1000 ha) accounted for gradually smaller amounts of total annual deforestation in the Brazilian Amazon, whereas the proportion of small clearings (<50 ha) nowadays represents more than 80% of annual deforestation. Despite the ubiquity of small clearings in fragmented Amazonian landscapes, most fragmentation research has focused on the effects of large-scale deforestation, leading to a poor understanding of the impacts of smaller barriers on Amazonian vertebrates. We capitalized on the periodical re-isolation of experimental forest fragments at the Biological Dynamics of Forest Fragments Project in the Central Amazon as a before-after-control-impact experiment to investigate the short-term effects of small clearings on bat assemblages. Over the course of three years we sampled six control sites in continuous forest, the interiors and edges of eight forest fragments as well as eight sites in the surrounding matrix. Sampling took place both before and after the experimental manipulation (clearing of a 100 m wide strip of regrowth around each fragment), resulting in ~4000 bat captures. Species were classified as old-growth specialists and habitat generalists according to their habitat affinities and a joint species distribution modeling framework was used to investigate the effect of fragment re-isolation on species occupancy. Following fragment re-isolation, species richness declined in all habitats other than fragment edges and, although responses were idiosyncratic, this decline was more pronounced for forest specialist than for generalist species. Additionally, fragment re-isolation led to a reduction in the similarity between assemblages in modified habitats (fragment interiors, edges and matrix) and continuous forest. Sampling of controls in continuous forest both prior to and after reisolation revealed that much of the variation in bat species occupancy between sampling periods did not arise from fragment re-isolation but rather reflected natural spatiotemporal variability. This emphasizes the need to sample experimental controls both before and after experimental manipulation and suggests caution in the interpretation of results from studies in which the effects of habitat transformations are assessed based solely on data collected using space-for-time substitution approaches

Joint species distribution modelling with the r-package Hmsc

Joint Species Distribution Modelling (JSDM) is becoming an increasingly popular statistical method for analysing data in community ecology. Hierarchical Modelling of Species Communities (HMSC) is a general and flexible framework for fitting JSDMs. HMSC allows the integration of community ecology data with data on environmental covariates, species traits, phylogenetic relationships and the spatio-temporal context of the study, providing predictive insights into community assembly processes from non-manipulative observational data of species communities. The full range of functionality of HMSC has remained restricted to Matlab users only. To make HMSC accessible to the wider community of ecologists, we introduce Hmsc 3.0, a user-friendly r implementation. We illustrate the use of the package by applying Hmsc 3.0 to a range of case studies on real and simulated data. The real data consist of bird counts in a spatio-temporally structured dataset, environmental covariates, species traits and phylogenetic relationships. Vignettes on simulated data involve single-species models, models of small communities, models of large species communities and models for large spatial data. We demonstrate the estimation of species responses to environmental covariates and how these depend on species traits, as well as the estimation of residual species associations. We demonstrate how to construct and fit models with different types of random effects, how to examine MCMC convergence, how to examine the explanatory and predictive powers of the models, how to assess parameter estimates and how to make predictions. We further demonstrate how Hmsc 3.0 can be applied to normally distributed data, count data and presence-absence data. The package, along with the extended vignettes, makes JSDM fitting and post-processing easily accessible to ecologists familiar with r.Peer reviewe

Direct and indirect effects of a pH gradient bring insights into the mechanisms driving prokaryotic community structures

Background: pH is frequently reported as the main driver for prokaryotic community structure in soils. However, pH changes are also linked to "spillover effects" on other chemical parameters (e.g., availability of Al, Fe, Mn, Zn, and Cu) and plant growth, but these indirect effects on the microbial communities are rarely investigated. Usually, pH also co-varies with some confounding factors, such as land use, soil management (e.g., tillage and chemical inputs), plant cover, and/or edapho-climatic conditions. So, a more comprehensive analysis of the direct and indirect effects of pH brings a better understanding of the mechanisms driving prokaryotic (archaeal and bacterial) community structures. Results: We evaluated an agricultural soil pH gradient (from 4 to 6, the typical range for tropical farms), in a liming gradient with confounding factors minimized, investigating relationships between prokaryotic communities (16S rRNA) and physical-chemical parameters (indirect effects). Correlations, hierarchical modeling of species communities (HMSC), and random forest (RF) modeling indicated that both direct and indirect effects of the pH gradient affected the prokaryotic communities. Some OTUs were more affected by the pH changes (e.g., some Actinobacteria), while others were more affected by the indirect pH effects (e.g., some Proteobacteria). HMSC detected a phylogenetic signal related to the effects. Both HMSC and RF indicated that the main indirect effect was the pH changes on the availability of some elements (e.g., Al, Fe, and Cu), and secondarily, effects on plant growth and nutrient cycling also affected the OTUs. Additionally, we found that some of the OTUs that responded to pH also correlated with CO2, CH4, and N2O greenhouse gas fluxes. Conclusions: Our results indicate that there are two distinct pH-related mechanisms driving prokaryotic community structures, the direct effect and "spillover effects" of pH (indirect effects). Moreover, the indirect effects are highly relevant for some OTUs and consequently for the community structure; therefore, it is a mechanism that should be further investigated in microbial ecology.Peer reviewe

Design matters : an evaluation of the impact of small man-made forest clearings on tropical bats using a before-after-control-impact design

In recent years, large clearings (>1000 ha) accounted for gradually smaller amounts of total annual deforestation in the Brazilian Amazon, whereas the proportion of small clearings (<50 ha) nowadays represents more than 80% of annual deforestation. Despite the ubiquity of small clearings in fragmented Amazonian landscapes, most fragmentation research has focused on the effects of large-scale deforestation, leading to a poor understanding of the impacts of smaller barriers on Amazonian vertebrates. We capitalized on the periodical re-isolation of experimental forest fragments at the Biological Dynamics of Forest Fragments Project in the Central Amazon as a before-after-control-impact experiment to investigate the short-term effects of small clearings on bat assemblages. Over the course of three years we sampled six control sites in continuous forest, the interiors and edges of eight forest fragments as well as eight sites in the surrounding matrix. Sampling took place both before and after the experimental manipulation (clearing of a 100 m wide strip of regrowth around each fragment), resulting in ~4000 bat captures. Species were classified as old-growth specialists and habitat generalists according to their habitat affinities and a joint species distribution modeling framework was used to investigate the effect of fragment re-isolation on species occupancy. Following fragment re-isolation, species richness declined in all habitats other than fragment edges and, although responses were idiosyncratic, this decline was more pronounced for forest specialist than for generalist species. Additionally, fragment re-isolation led to a reduction in the similarity between assemblages in modified habitats (fragment interiors, edges and matrix) and continuous forest. Sampling of controls in continuous forest both prior to and after reisolation revealed that much of the variation in bat species occupancy between sampling periods did not arise from fragment re-isolation but rather reflected natural spatiotemporal variability. This emphasizes the need to sample experimental controls both before and after experimental manipulation and suggests caution in the interpretation of results from studies in which the effects of habitat transformations are assessed based solely on data collected using space-for-time substitution approaches

Invasion rate of deer ked depends on spatiotemporal variation in host density

Invasive parasites are of great global concern. Understanding the factors influencing the spread of invading pest species is a first step in developing effective countermeasures. Growing empirical evidence suggests that spread rates are essentially influenced by spatiotemporal dynamics of host-parasite interactions, yet approaches modelling spread rate have typically assumed static environmental conditions. We analysed invasion history of the deer ked (Lipoptena cervi) in Finland with a diffusion-reaction model, which assumed either the movement rate, the population growth rate, or both rates may depend on spatial and temporal distribution of moose (Alces alces), the main host of deer ked. We fitted the model to the data in a Bayesian framework, and used the Bayesian information criterion to show that accounting for the variation in local moose density improved the model's ability to describe the pattern of the invasion. The highest ranked model predicted higher movement rate and growth rate of deer ked with increasing moose density. Our results suggest that the historic increase in host density has facilitated the spread of the deer ked. Our approach illustrates how information about the ecology of an invasive species can be extracted from the spatial pattern of spread even with rather limited dat