Francesco de Bello’s research while affiliated with Desertification Research Centre and other places

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Publications (255)


Estimating dark diversity and related biodiversity metrics in ecological communities
a, Data included a local study site where certain species were present, but many species sampled elsewhere in the region were absent. To estimate the probability that a species that is absent from the site but present in the region belongs to the dark diversity of the site, we used information about species co-occurrences at other sites in the region. b, We calculated an indicator matrix in which each present species indicated the ecological suitability of each absent species for the study site. We compared the observed number of co-occurrences with the number of co-occurrences expected at random (according to the hypergeometric distribution) and standardized the difference using the standard deviation from the hypergeometric distribution. c, By averaging across all observed species, each absent species was assigned a probability of belonging to the dark diversity for the study site. Consequently, the dark diversity was a fuzzy set to which species belonged to varying degrees. d, Several biodiversity metrics were characterized for each site in the region. Alpha diversity was the number of species recorded at the site, and gamma diversity was the total number of species recorded in a region. The size of dark diversity was estimated as the sum of the probabilities of absent species belonging to the dark diversity of the study site. Alpha and dark diversity together formed the site-specific species pool, and gamma diversity not falling into this category was considered the unsuitable part of gamma diversity; that is, belonging to the species pools of other sites. We investigated the percentage of the species pool that was present among the alpha diversity (community completeness) and the turnover of species pools in the region, expressed as the percentage of gamma diversity that was unsuitable for the study site (beta diversity).
Plant diversity in natural vegetation in relation to human effects in the surrounding regions
a, Relationship between community completeness in natural vegetation and the human footprint index in the surrounding area, defined by a radius of 300 km. The prediction line from a multiple linear regression model is shown with the 95% confidence intervals. Note that community completeness values on the y axis are back-transformed from the logit scale. The symbol tones indicate forest cover (0–100%). R² value of the model and two-tailed P value of the relationship are shown; n = 116 regions. The distribution of community completeness is shown in the histogram on the right (median, 25%). b, Left, model summaries linking community completeness to the human footprint index and its components across spatial scales. Human influence was averaged over various spatial scales around the study regions (radii 10 km, 50 km, 100 km, 200 km, 300 km and 400 km), and the respective models were compared using the Akaike information criterion (AIC). Filled symbols indicate significant relationships (P < 0.05), and the large symbol indicates the set of best significant models (ΔAIC < 2). Right, from the best model (the smallest scale at which ΔAIC < 2), the effect of the human footprint index or one of its components is shown as a standardized coefficient (dot) with a 95% confidence interval (CI; line); n = 116 regions. Filled symbols and bold confidence interval lines indicate significant effects. c, Map of sampling regions, with community completeness indicated by symbol size and the underlying map showing the global variation in the human footprint index³⁴ (the highest value within each grid cell of around 0.25° × 0.25°). The inset shows part of Europe containing a large number of study regions. Triangles indicate regions in which only woody species were sampled. Symbol tones indicate the percentage of forests in regions.
Global impoverishment of natural vegetation revealed by dark diversity
  • Article
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April 2025

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3,151 Reads

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3 Citations

Nature

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Anthropogenic biodiversity decline threatens the functioning of ecosystems and the many benefits they provide to humanity¹. As well as causing species losses in directly affected locations, human influence might also reduce biodiversity in relatively unmodified vegetation if far-reaching anthropogenic effects trigger local extinctions and hinder recolonization. Here we show that local plant diversity is globally negatively related to the level of anthropogenic activity in the surrounding region. Impoverishment of natural vegetation was evident only when we considered community completeness: the proportion of all suitable species in the region that are present at a site. To estimate community completeness, we compared the number of recorded species with the dark diversity—ecologically suitable species that are absent from a site but present in the surrounding region². In the sampled regions with a minimal human footprint index, an average of 35% of suitable plant species were present locally, compared with less than 20% in highly affected regions. Besides having the potential to uncover overlooked threats to biodiversity, dark diversity also provides guidance for nature conservation. Species in the dark diversity remain regionally present, and their local populations might be restored through measures that improve connectivity between natural vegetation fragments and reduce threats to population persistence.

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Handbook of protocols for standardized measurements of floral traits for pollinators in temperate communities

March 2025

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333 Reads

Floral traits describe organs or structures directly related to plant reproduction, and they are essential to understanding plant–pollinator interactions, notably for conservation purposes. The growth of plant trait‐based approaches lies in the availability of data shared by the international research community on dedicated platforms, as well as in protocols compiled in handbooks on how to measure these traits in a standardized way. Floral traits are important pieces that are missing from these handbooks, likely due to the complexity of flower morphology. Here, we present a handbook of standardized protocols dedicated to floral traits that can be applied to a wide set of temperate plant species to quantify these traits at the scale of plant communities. The 24 floral traits are grouped into three categories: visual and olfactory cues, accessibility and resources. We also provide four additional features related to flower abundance and phenology that we recommend measuring to scale up individual species' trait values to overall plant communities. By collecting these floral traits in a standardized way, we promote applications in the context of community ecology to predict the diversity of pollinator communities, identify the effects of environmental changes and study plant–pollinator networks.



Functional traits mediate the effect of land use on drivers of community stability within and across trophic levels

January 2025

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572 Reads

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1 Citation

Science Advances

Understanding how land use affects temporal stability is crucial to preserve biodiversity and ecosystem functions. Yet, the mechanistic links between land-use intensity and stability-driving mechanisms remain unclear, with functional traits likely playing a key role. Using 13 years of data from 300 sites in Germany, we tested whether and how trait-based community features mediate the effect of land-use intensity on acknowledged stability drivers (compensatory dynamics, portfolio effect, and dominant species variability), within and across plant and arthropod communities. Trait-based plant features, especially the prevalence of acquisitive strategies along the leaf-economics spectrum, were the main land-use intensity mediators within and across taxonomic and trophic levels, consistently influencing dominant species variability. Functional diversity also mediated land-use intensity effects but played a lesser role. Our analysis discloses trait-based community features as key mediators of land-use effects on stability drivers, emphasizing the need to consider multi-trophic functional interactions to better understand complex ecosystem dynamics.


There is no single trait–environment relationship

November 2024

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221 Reads

A popular approach assessing trait–environment relationships is the Community Weighted Means (CWMs) method, which evaluates changes in communities' average trait values along gradients. Recently, the use of CWM as response in general linear models have been criticized for inflated Type I errors. In some scenarios of compositional turnover along a gradient, CWM tests can provide significant results even for randomly generated traits. Null models have been proposed to correct for such effects. However, the results of analyses relating traits to the environment must be interpreted within the framework of the accepted dichotomy between species-level (observations are species) vs. community-level (observations are community parameters) analyses, like with CWM. Within this framework, Type I error rate with CWM tests should not be a priori considered inflated when community-level analysis is used to identify changes in community trait structure. On the other hand, identifying specific traits that cause species turnover, and species adaptation, is more appropriate with methods approaching the species-level analyses. Between these families of analyses, a great variety of methods exist that test different trait–environment relationships, each with different null hypotheses and ecological questions. There is no single trait–environment relationship. Using a spectrum of methods provides a comprehensive picture of the diversity of trait–environment relationships.


The effect of neighbor species' phylogenetic and trait difference on tree growth in subtropical forests

August 2024

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137 Reads

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1 Citation

Journal of Vegetation Science

Questions To comprehensively understand ecological dynamics within a forest ecosystem, it is vital to explore how surrounding trees influence the growth of individual trees in a community. This study investigates the importance of biotic interactions on tree growth by examining several metrics of competitive interactions and community structure and considering three classes of intrinsic growth rates among the focal individuals: slower, intermediate, and faster‐growing trees. We also separated the focal trees based on their canopy position. Location Brazilian subtropical forests. Methods We assessed various factors related to the focal trees and their neighbors, including differences in traits, neighborhood crowding, phylogenetic distance, and overall trait composition within the community. We then ran linear mixed‐effects models to test how these different metrics influenced the growth rates of the focal trees. Results Our results indicate that phylogenetic distance is linked to higher growth. Specific leaf area (SLA), leaf area (LA), and wood density (WD) are significantly related to tree growth. Trees surrounded by neighbors with higher SLA than themselves grow better, particularly smaller trees. Similarly, taller trees with smaller LA than their neighbors grow better. Trees in the intermediary growth class grow better when they have higher WD than their neighbors. Conversely, smaller trees benefit from greater WD difference between the focal trees and their neighbors, while height difference negatively impacts faster‐growing trees. Moreover, communities with higher SLA and WD positively impact the growth of faster‐growing trees. Conclusions We conclude that the interactions between trees are mediated by their ecological differences, but the performance and responses to surrounding competitors vary along with their grow class and position within a community. This study has revealed that the tree's intrinsic growth rate mediates the effect of traits and phylogeny of surrounding trees on individual tree growth.


ReSurveyEurope: A database of resurveyed vegetation plots in Europe

April 2024

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1,164 Reads

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15 Citations

Journal of Vegetation Science

Aims: We introduce ReSurveyEurope — a new data source of resurveyed vegetation plots in Europe, compiled by a collaborative network of vegetation scientists. We describe the scope of this initiative, provide an overview of currently available data, governance, data contribution rules, and accessibility. In addition, we outline further steps, including potential research questions. Results: ReSurveyEurope includes resurveyed vegetation plots from all habitats. Version 1.0 of ReSurveyEurope contains 283,135 observations (i.e., individual surveys of each plot) from 79,190 plots sampled in 449 independent resurvey projects. Of these, 62,139 (78%) are permanent plots, that is, marked in situ, or located with GPS, which allow for high spatial accuracy in resurvey. The remaining 17,051 (22%) plots are from studies in which plots from the initial survey could not be exactly relocated. Four data sets, which together account for 28,470 (36%) plots, provide only presence/absence information on plant species, while the remaining 50,720 (64%) plots contain abundance information (e.g., percentage cover or cover–abundance classes such as variants of the Braun-Blanquet scale). The oldest plots were sampled in 1911 in the Swiss Alps, while most plots were sampled between 1950 and 2020. Conclusions: ReSurveyEurope is a new resource to address a wide range of research questions on fine-scale changes in European vegetation. The initiative is devoted to an inclusive and transparent governance and data usage approach, based on slightly adapted rules of the well-established European Vegetation Archive (EVA). ReSurveyEurope data are ready for use, and proposals for analyses of the data set can be submitted at any time to the coordinators. Still, further data contributions are highly welcome.



Figure A1, Appendix A1 for supplementary information on the PCA). Positive values of the slow-fast continuum were associated with plant communities dominated by species with acquisitive strategies (fast-growing species), whereas negative values represented plant communities characterised by species with conservative strategies (slow-growing species). Note that, to compute community-weighted means, trait values were weighted by species cover, which we measured (on a percentage scale) in the same 4 m × 4 m sub-quadrats
Biodiversity promotes resistance but dominant species shape recovery of grasslands under extreme drought

March 2024

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464 Reads

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15 Citations

1. How biodiversity underpins ecosystem resistance (i.e. ability to withstand environmental perturbations) and recovery (i.e. ability to return to a pre-perturbation state), and thus, stability under extreme climatic events is a timely question in ecology. To date, most studies have focussed on the role of taxonomic diversity, neglecting how community functional composition and diversity beget stability under exceptional climatic conditions. In addition, land use potentially modulates how biodiversity and ecosystem functions respond to extreme climatic conditions. 2. Using an 11-year time-series of plant biomass from 150 permanent grassland plots spanning a gradient of land-use intensity, we examined how taxonomic and functional components of biodiversity affected resistance and recovery of biomass under extreme drought. 3. The association between biodiversity, land use and biomass varied across years, especially in the driest years. Species-rich or functionally diverse communities (associated with low land-use intensity) buffered extreme droughts better, while species-poor communities or those dominated by fast-growing species (associated with high land-use intensity) had higher recovery capabilities after a moderate-to-extreme drought. 4. Synthesis. Our results show that plant community functional and taxonomic components determine grasslands resistance and recovery under moderate-to-extreme drought. In turn, this points to the importance of designing landscapes with both extensively and intensively managed grasslands. Functionally or taxonomically rich communities (favoured under low land-use intensity) would preserve biomass under extreme droughts, whereas species-poor or fast-growing communities (favoured by high land-use intensity) would restore biomass after extreme droughts.


Figure 1. (a) The temporal stability of communities is simultaneously driven by four distinct components related to species characteristics and the community structure. These four components are conceptually represented in the central panels where the same community fluctuation (bold line) results from the fluctuations of different species (colored dotted lines) representing (a.1) globally stable species, the stabilizing role of (a.2) a dominant species, or of compensatory dynamics due to (a.3) the asynchrony of species or (a.4) the averaging effect (associated with high diversity). (b) Our unified framework allows us to disentangle all four components of stability occurring simultaneously in a community and represent them on a simple graph that describes their relative importance (see “General framework” section for details on the indices and Box 1 for the main definitions of the terms proposed in the framework). Briefly, CVe is the variability of a species of mean abundance (related to panel a.1) which is stabilized by dominance (Δ, related to panel a.2), asynchrony (ψ, related to panel a.3), and averaging (ω, related to panel a.4) effects to give the community variability CVcom (reciprocal of stability). (CV) ̃ is the abundance-weighted mean of species CVi, and CVa is the expected CVcom if the community was stabilized by species asynchrony only
Figure 2. Temporal fluctuation of four simulated communities representing different scenarios of species richness, dominance, mean-variance scaling, and asynchrony. For each scenario, (1) the selected species richness and dominance are represented by the rank-abundance plot, (2) the log-log relationship between species means and variances shows the comparison between the selected scaling coefficient (colored dotted line), and 2 (solid line) and (3) the annual fluctuations of all species abundances show the selected synchrony level (see main text for the exact values). Four panels show the partitioning of the corresponding community coefficient of variation (CVcom) into the variability of the average species (CVe) and three stabilizing effects: the dominance (Δ), asynchrony (ψ), and averaging (ω) effects. (CV) ̃ is the abundance-weighted mean of species CVi and CVa is the expected CVcom if the community was stabilized by species asynchrony only. Y-axes of these panels are log-transformed so the strength of each stabilizing effect is represented by the slope between two successive points: a steep negative slope means a high stabilization. Finally, the ternary plot in the center shows the relative contributions of the three stabilizing effects to the total stabilization (τ) for the four communities (symbol of the corresponding color). Tick orientation shows how to read the values. Note that for each of the scenarios, despite their different prominent stabilizing effects, CVcom, CVe, and the overall stabilization are kept the same.
Figure 3. Effect of (a) fertilization and (b) dominant removal on the temporal coefficient of variation (CVcom) of a grassland community. The coefficient of variation of the average species (CVe) is stabilized by the dominance (Δ), asynchrony (ψ), and averaging (ω) effects. (CV) ̃ is the abundance-weighted mean of species CVi and CVa is the expected CVcom if the community was stabilized by species asynchrony only. The total stabilizations (τ) are given in the panel legends. Y-axes are log-transformed so the strength of each stabilizing effect is represented by the slope between two successive points (the corresponding values are given next to each line): a steep negative slope means a high stabilization. Asynchrony values corresponding to a prevalent positive or negative synchrony between species are represented by a (+) or (-), respectively.
Figure 4. Relative contributions of dominance, asynchrony, and averaging effects on the total stabilization of grassland communities in control plots (red symbol), fertilized plots (orange symbol), and plots where the most dominant species has been removed (blue symbol). The axes represent the contribution of each stabilizing effect to the total stabilization, calculated as the ratios between the corresponding log-transformed index (log(Δ), log(ψ) or, log(ω)) and the log-transformed total stabilization (log(τ)). Tick orientation shows how to read the values, e.g., for the orange symbol, the contribution of dominance, asynchrony, and averaging effects to the total stabilization are roughly 20 %, 30 %, and 50 %, respectively.
A unified framework for partitioning the drivers of stability of ecological communities

March 2024

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1,130 Reads

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9 Citations

Aim Identifying the drivers of ecological stability is critical for ensuring the maintenance of ecosystem functioning and services, particularly in a changing world. Different ecological mechanisms by which biological communities stabilize ecosystem functions (i.e. “stabilizing effects”) have been proposed, yet with various theoretical expectations and debated conclusions. Here we propose a unified framework that aims at reconciling, and combining, different approaches to reliably test the strength of three stabilizing effects on the temporal constancy of ecosystem functions: the effects of (a) dominant species, (b) species asynchrony, and (c) diversity. Innovation Compared to existing developments the approach allows, for the first time, disentangling these three stabilizing effects at the level of individual communities. So far this was not possible, and conclusions depended on indirect tests and comparative analyses across communities. We also propose a graphical representation of the relative contributions of the three stabilizing effects on a ternary plot, allowing us to easily compare communities sampled in various ecological contexts in a standardized space. Main conclusions Our study answers the current need for a unified framework to link theoretical concepts on the temporal stability of ecological communities to data analysis. The present development promises flexible tests for a deeper understanding of the ecological stabilization of biodiversity and the relative importance of its components.


Citations (81)


... The success of this approach is underscored by the creation of centers for ecological synthesis, such as the National Center for Ecological Analysis and Synthesis (NCEAS) in the United States or the Synthesis Centre for Biodiversity Sciences in Germany (sDiv). In parallel, other research groups have developed distributed ecological networks of experiments and/or observations, for example, NutNet or DarkDivNet (Borer et al. 2014;Pärtel et al. 2025). In these projects, researchers focus on different locations, but they all implement standardized methods to yield geographically comparative data. ...

Reference:

Editorial : Biotropica Announces a New Paper Category: Synthesis
Global impoverishment of natural vegetation revealed by dark diversity

Nature

... In the future, including plant functional trait-based community features (i.e. trait composition and diversity) could provide further insights into how different plant communities interact with environmental conditions and contribute to ecosystem functions and respond to environmental changes (Galán Díaz et al., 2022;Sperandii et al., 2025). ...

Functional traits mediate the effect of land use on drivers of community stability within and across trophic levels
  • Citing Article
  • January 2025

Science Advances

... ( Figure 5). Likewise, data from repeatedly sampled plots (permanent or quasi-permanent plots) could more beneficially be contributed to ReSurveyEurope (Knollová et al. 2024) ( Figure 5). Note that despite belonging to the Nordic-Baltic region, grassland plots s.l. ...

ReSurveyEurope: A database of resurveyed vegetation plots in Europe

Journal of Vegetation Science

... Field-based studies, while valuable for localized and detailed insights, are inherently constrained by logistical challenges, including the difficulty of accessing remote or rugged terrains and the time-consuming nature of data collection [23]. Moreover, they lack the capacity to capture rapid or large-scale changes across diverse landscapes, making it difficult to establish comprehensive and timely assessments [24,25]. There is a growing need for advanced tools that can provide automated, scalable, and reproducible assessments of landscape changes. ...

ReSurveyEurope: A database of resurveyed vegetation plots in Europe

Journal of Vegetation Science

... (1)) at both pixel and tree level. The suitability of Rao's index for calculating different aspects of heterogeneity from remote sensing data had been previously validated in other studies (Perrone et al., 2024;Torresani et al., 2024a;Michele et al., 2018;Tamburlin et al., 2021;Torresani et al., 2024b). In contrast, its application for detecting HH at tree level was explored for the first time in this study. ...

“Flower power”: How flowering affects spectral diversity metrics and their relationship with plant diversity

Ecological Informatics

... The high dynamics in environmental parameters suggested that the Hong Kong waters were strongly inf luenced by anthropogenic impacts or natural disturbances (Fig. S1). Although previous studies indicated that core taxa tend to exhibit relatively higher sensitivity and resilience in response to multiple environmental stressors [34,42], diversity of three core groups displayed distinct seasonal and geographical patterns, ref lecting their unique ecological strategies and potential adaptive mechanisms (Fig. 2). Despite the high local Shannon diversity, the core autotrophic communities demonstrate a high stability in α-diversity, providing fundamental resources for the community. ...

Biodiversity promotes resistance but dominant species shape recovery of grasslands under extreme drought

... This suggests that higher species asynchrony buffers the effects of environmental fluctuations on species, promotes compensatory species dynamics, and reduces temporal variability in community function, thereby improving ecosystem stability (Loreau and de Mazancourt, 2008;Zhang et al., 2016;Xu et al., 2021;Li et al., 2021). Significant positive correlations between dominant species stability and ecosystem stability were found in 80 % of the datasets (Fig. 4c, d), suggesting that dominant species are more adaptive to the environment and fluctuate less from year to year, thus maintaining the stability of ecosystem functioning (Grman et al., 2010;Segrestin et al., 2024). The positive species richness-ecosystem stability correlation was not strong, and we only found significant positive correlations in 33 % of the datasets (Fig. 4e, f). ...

A unified framework for partitioning the drivers of stability of ecological communities

... We investigated whether the Fundão tailings dam failure changed (1) ant species richness, ant foraging activity and species composition; (2) the phylogenetic structure of the ant community; and (3) the ecosystem function of seed removal by ants. Specifically, we hypothesize that this disturbance will cause a phylogenetic impoverishment (decrease in the phylogenetic richness) as a consequence of species loss (Ribeiro et al. 2016), and that it will lead to phylogenetic clustering, a pattern often observed in ant communities following ecological disturbance (Vasconcelos et al. 2024;Pérez-Toledo et al. 2024, Hoenle et al. 2024Arnan et al. 2018). Additionally, we expect that this disturbance will impact ants ecosystem function of seed dispersal. ...

Forest disturbance increases functional diversity but decreases phylogenetic diversity of an arboreal tropical ant community

... The selection of traits was determined by (i) the a priori assumption about their sensitivity to and relevance with climate and land use spatiotemporal changes (e.g., [45,46]), (ii) their prior utilization in assessing the functional diversity of and processes of community assembly in amphibians (see Tsianou and Kallimanis [47][48][49]), and (iii) the extent to which speciesspecific trait information was available. The importance of intraspecific trait variability (ITV)-including variability between cryptic species-in functional diversity patterns has been gaining recognition lately (e.g., [50][51][52][53]). However, we were not able to address the potential influence of ITV due to the lack of the required population-level trait data across the entire extent of occurrence for each analyzed species. ...

Integrating intraspecific trait variability in functional diversity: an overview of methods and a guide for ecologists

... If redundant species are similar in their ecosystem functions but differ in their environmental preferences, redundancy may enhance community functioning and stability after environmental perturbations. For example, if a nitrogen-fixing species is lost due to environmental perturbations, its role can be partially compensated by the presence of other nitrogen fixers, which are more tolerant to perturbation (Fischer and de Bello 2023). Based on the assumption that ecological differences between species are related to their functional dissimilarities, such that functionally similar species have similar ecological roles, Leinster and Cobbold (2012) defined the ordinariness of species i as the abundance of all species that are functionally similar to i (including i itself): ...

On the uniqueness of functional redundancy

npj Biodiversity