Carlos P. Carmona

University of Tartu · Department of Botany

Islands are biodiversity hotspots that host unique assemblages. However, a substantial proportion of island species are threatened and their long‐term survival is uncertain. Identifying and preserving vulnerable species has become a priority, but it is also essential to combine this information with other facets of biodiversity like functional dive...

Mammalian life history strategies can be characterised by a few axes of variation, conforming a space where species are positioned based on the life history strategies favoured in the environment they exploit. Yet, we still lack global descriptions of the diversity of realised mammalian life history and how this diversity is shaped by the environme...

Dark diversity includes ecologically suitable species currently absent in a site, albeit theoretically able to arrive from the surrounding region. Various methods can estimate the likelihood that an absent species is in the dark diversity of a site. Recent developments in estimation of dark diversity have advanced the field, yet uncertainty on meth...

Differences within species (Intraspecific trait variation - ITV) contribute substantially to overall trait variability and environmental harshness can reduce among-species variation. While aboveground traits have received considerable attention, knowledge about ITV in fine-root traits and how it differs from ITV in aboveground traits remains limite...

Aim Combining different biodiversity dimensions can reveal new diversity patterns disclosing the relative roles of historical, environmental and anthropogenic factors in shaping global seed plant diversity. Location Global. Time period Present. Major taxa studied Vascular plants. Methods We collated a database encompassing taxonomic (249,000 sp...

Theory suggests that intraspecific trait variability may promote species coexistence when competitively inferior species have higher intraspecific trait variability than their superior competitors. Here, we provide empirical evidence for this phenomenon in tree seedlings. We evaluated intraspecific variability and plastic response of ten traits in...

Question Remote sensing is a fundamental tool to monitor biodiversity and spectral diversity may represent a proxy for different biodiversity facets such as taxonomic (TD) and functional diversity (FD). We used fine‐resolution multispectral imagery to explore spectral diversity (SD) patterns across spatial scales (i.e., plot, transect, area), and a...

Aim Functional trait space analyses are pivotal to describe and compare organisms' functional diversity across the tree of life. Yet, there is no single application that streamlines the many sometimes‐troublesome steps needed to build and analyse functional trait spaces. Innovation To fill this gap, we propose funspace, an R package to easily hand...

The Kunming–Montreal Global Biodiversity Framework of the Convention on Biological Diversity calls for an expansion of the current protected areas (PAs) to cover at least 30% of global land and water areas by 2030 (i.e., the 30×30 target). Efficient spatial planning for PA expansion is an urgent need for global conservation practice. A spatial prio...

Human activities have altered the species composition of assemblages through introductions and extinctions, but it remains unclear how those changes can affect the different facets of biodiversity. Here we assessed the impact of changes in species composition on taxonomic, functional, and phylogenetic diversity across 281 bird assemblages worldwide...

Question Trait‐based ecology is built on the notion that traits impact individual performance, but trait–growth relationships have rarely been tested considering the intraspecific variation, trait interactions, and potential nonlinearity in these relationships. Location Seven tropical dry forests throughout Colombia. Methods We examined trait–gro...

The variability in traits within species (intraspecific trait variability; ITV) has attracted an increased interest in functional ecology, as it can profoundly influence the detection of functional trait patterns, calculation of functional diversity (FD), and assessments of ecosystem functioning. This renewed focus stems from the recognition that s...

Biodiversity monitoring is constrained by cost- and labour-intensive field sampling methods. Increasing evidence suggests that remotely sensed spectral diversity (SD) is linked to plant diversity, holding promise for monitoring applications. However, studies testing such a relationship reported conflicting findings, especially in challenging ecosys...

Aim Trade‐offs among tolerances to different abiotic stressors limit polytolerance in woody plants. However, the general trait syndromes that underlie large‐scale tolerance patterns of woody plants remain controversial. Here, we tested if the leading trait dimensions that define the global spectrum of plant form and function capture the underlying...

Human activities have altered the species composition of assemblages through introductions and extinctions, but it remains unclear how those changes can affect the different facets of biodiversity. Here we assessed the impact of changes in species composition on taxonomic, functional, and phylogenetic diversity across 281 bird assemblages worldwide...

Characterising changes in functional diversity at large spatial scales provides insight into the impact of human activity on ecosystem structure and function. However, the approach is often based on trait data sets that are incomplete and unrepresentative, with uncertain impacts on functional diversity estimates. To address this knowledge gap, we s...

Mammalian life history strategies can be characterized by a few axes of variation, which conform a space where species are positioned according to which life history strategies are favoured in the environment they exploit. Yet, we still lack global descriptions of the diversity of realized mammalian life history and of how this diversity is shaped...

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetati...

In the accompanying Comment, Weigelt et al. (2023) claim that the different conclusions of two studies—Weigelt et al. (2021) and Carmona et al. (2021)—presenting global integrations of aboveground and belowground plant traits are due to methodological issues in Carmona et al. Specifically, Weigelt et al. concluded that leaf and fine-root traits are...

This poster presents the evaluation of functional diversity indices as early-warning indicators for rangeland degradation in semi-arid savannas in Namibia. We present our study set-up, the data collection and variables, that were used to calculate functional diversity indices. Based on that we show analyses regarding threshold behaviour and the fea...

Although functional traits are defined based on their impact on demographic parameters, trait-demography relationships are often reported as weak. These weak relationships might be due to disregarding trait interactions and environmental contexts, which should modulate species trait-demography relationships. We applied different models, including b...

The Anthropocene is tightly associated with a drastic loss of species worldwide and the disappearance of their key ecosystem functions. The orders Testudines (turtles and tortoises) and Crocodilia (crocodiles, alligators, and gharials) contain numerous threatened, long-lived species for which the functional diversity and potential erosion by anthro...

1. Functional trait space analyses are pivotal to define species' ecological strategies across the tree of life. Yet, there is no single application that streamlines the many sometimes-troublesome steps needed to build and analyze functional trait spaces. 2. To fill this gap, we propose funspace, an R package to easily handle bivariate and multivar...

Human activities have altered the composition of species assemblages through the introduction of non-native species and the extinction of threatened species. However, it remains unclear whether non-native species can compensate for the loss of threatened species and thus maintain ecosystem functioning. Here we tested whether non-natives are functio...

Remote sensing is a fundamental tool to monitor biodiversity over large spatial extents. However, it is still not clear whether spectral diversity (SD - variation of spectral response across a set of pixels) may represent a fast and reliable proxy for different biodiversity facets such as taxonomic (TD) and functional diversity (FD) across differen...

Despite evidence of declining biosphere integrity, we currently lack understanding of how the functional diversity associated with changes in abundance among ecological communities has varied over time and before widespread human disturbances. We combine morphological, ecological, and life-history trait data for >260 extant bird species with genomi...

Global change drivers (e.g. climate and land use) affect the species and functional traits observed in a local site but also its dark diversity—the set of species and traits locally suitable but absent. Dark diversity links regional and local scales and, over time, reveals taxa under expansion lags by depicting the potential biodiversity that remai...

1. Characterizing changes in trait diversity at large spatial scales provides insight into the impact of human activity on ecosystem structure and function. However, the approach is often based on trait datasets that are incomplete and unrepresentative, with uncertain impacts on trait diversity estimates. 2. To address this knowledge gap, we simula...

While biodiversity is expected to enhance multiple ecosystem functions (EFs), the different roles of multiple biodiversity dimensions remain difficult to disentangle without carefully designed experiments. We sowed plant communities with independent levels of functional (FD) and phylogenetic diversities (PD), combined with different levels of ferti...

Understory plants contribute to >80% of the understory forest diversity and are susceptible to climatic changes. We analysed long-term monitoring and climate data to determine: (a) what are the patterns of functional diversity (FD) through time and across forests, (b) does climate explain these patterns, and (c) does above and belowground FD respon...

Aim Analysis of the shifts in the climatic space for invasive species between native and introduced areas is a powerful tool for understanding their distribution patterns and the factors influencing their spread into new areas. In this study, we explored the shift in climatic distribution between ranges for invasive woody legumes and how the functi...

In trait‐based ecology, phenotypic variation (PVar) is often quantified with measures expressing average differences between populations standardized in the range 0–1. However, these measures disregard the within‐population trait variability. In addition, some of them cannot be partitioned between populations. These aspects can either alter their i...

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species’ ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, specially across different ecosystems. To address this, we compiled a global collection of long-term p...

Drought is expected to increase in future climate scenarios. Although responses to drought of individual functional traits are relatively well‐known, simultaneous changes across multiple traits in response to water scarcity remain poorly understood despite its importance to understand alternative strategies to resist drought. We grew 52 herbaceous...

The Anthropocene is tightly associated with a drastic loss of species worldwide, and consequently, with the disappearance of key ecosystem functions. The ongoing reduction in species functionality is driven by global and local threats. The orders Testudines (turtles and tortoises) and Crocodilia (crocodilians and alligators) contain numerous threat...

Purpose Plant species diversity is expected to affect multiple ecosystem functions, such as soil nitrogen (N) availability. However, this effect may be related to the ecological differentiation between coexisting species, often expressed as either functional diversity (FD; diversity in traits) or phylogenetic diversity (PD; diversity in phylogeneti...

Phenotypic plasticity is essential for organisms to adapt to local ecological conditions. It is expected that mutualistic interactions, such as arbuscular mycorrhizal (AM) symbiosis, mediate plant phenotypic plasticity, although it is not clear to what extent this plasticity may be heritable (i.e. transgenerational plasticity). We tested for plant...

Trait and functional trait approaches have revolutionized ecology improving our understanding of community assembly, species coexistence, and biodiversity loss. Focusing on traits promotes comparability across spatial and organizational scales, but terms must be used consistently. While several papers have offered definitions, it remains unclear ho...

In trait-based ecology, phenotypic variation (PVar) is often quantified with measures that express average differences between populations standardized in the range 0-1. A major problem with these measures is that they disregard the within-population trait variability. In addition, most of these measures cannot be decomposed across scales. This can...

Installing patches of flowering plants is a commonly used strategy to enhance refuge and food resources for pollinators in intensive agricultural landscapes. Here, we evaluated how floral strips and semi-natural habitats impact the taxonomic and functional diversity of wild bees in intensively farmed sunflower fields. Pan traps were used to assess...

Plant traits determine how individual plants cope with heterogeneous environments. Despite large variability in individual traits, trait coordination and trade-offs1,2 result in some trait combinations being much more widespread than others, as revealed in the global spectrum of plant form and function (GSPFF3) and the root economics space (RES4) f...

Although species with larger body size and slow pace of life have a higher risk of extinction at a global scale, it is unclear whether this global trend will be consistent across biogeographic realms. Here we measure the functional diversity of terrestrial and freshwater vertebrates in the six terrestrial biogeographic realms and predict their futu...

The use of functional diversity analyses in ecology has grown exponentially over the past two decades, broadening our understanding of biological diversity and its change across space and time. Virtually all ecological sub‐disciplines recognise the critical value of looking at species and communities from a functional perspective, and this has led...

The exploitation of non-timber forest products (NTFPs) has been encouraged in order to reconcile economic uses, biodiversity conservation and the provision of ecosystem services. In this paper we investigate how increments on the açaí palm density (Euterpe oleracea) via forest management change tree assemblages across 47 plots in the Amazon estuari...

We are in the midst of a sixth mass extinction but little is known about the global patterns of biodiversity when accounting for taxonomic, phylogenetic and functional information. Here, we present the first integrated analysis of global variation in taxonomic, functional diversity and phylogenetic diversity of more than 17,000 tetrapod species (te...

Mountains provide a timely opportunity to examine the potential effects of climate change on biodiversity. However, nature conservation in mountain areas have mostly focused on the observed part of biodiversity, not revealing the suitable but absent species—dark diversity. Dark diversity allows calculating the community completeness, indicating whe...

Aims The link between spectral diversity and in-situ plant biodiversity is one promising approach for using remote sensing for biodiversity assessment. Nevertheless, there is little evidence to whether this link is maintained at fine scales, as well as to how it is influenced by vegetation’s vertical complexity. Here we test, at community level in...

1. We address associations of taxonomic diversity (TD), functional diversity (FD), and phylogenetic diversity (PD) of ant assemblages with gradients of elevation to assess whether energetic limitations or deterministic or stochastic niche-building processes are more relevant to the assembly of communities. 2. We sampled ant assemblages using pitfal...

Background and aims: The observed positive diversity effect on ecosystem functioning has been scarcely assessed in terms of intraspecific trait variability within populations. Intraspecific phenotypic variability could stem both from underlying genetic diversity and plasticity in response to environmental cues. The latter might derive from modific...

Handbook of Trait-Based Ecology - From Theory to R Tools Functional ecology is the branch of ecology that focuses on various functions that species play in the community or ecosystem in which they occur. This accessible guide offers the main concepts and tools in trait-based ecology, and their tricks, covering different trophic levels and organism...

Although one-quarter of plant and vertebrate species are threatened with extinction, little is known about the potential effect of extinctions on the global diversity of ecological strategies. Using trait and phylogenetic information for more than 75,000 species of vascular plants, mammals, birds, reptiles, amphibians, and freshwater fish, we chara...

Questions What are the functional trade‐offs of vascular plant species in global alpine ecosystems? How is functional variation related to vegetation zones, climatic groups and biogeographic realms? What is the relative contribution of macroclimate and evolutionary history in shaping the functional variation of alpine plant communities? Location G...

Functional diversity assessments are crucial and increasingly used for understanding ecological processes and managing ecosystems. The functional diversity of a community is assessed by sampling traits at one or more scales (individuals, populations and species) and calculating a summary index of the variation in trait values. However, it remains u...

The use of functional analyses in ecology has grown exponentially over the past two decades, broadening our understanding of biological diversity and its change across space and time. Virtually all ecological sub-disciplines recognize the critical value of looking at species and communities from a functional perspective, and this has led to a proli...

Extreme drought events have negative effects on forest diversity and functioning. At the species level, however, these effects are still unclear, as species vary in their response to drought through specific functional trait combinations. We used long-term demographic records of 21,821 trees and extensive databases of traits to understand the respo...

Phenotypic plasticity, within and across generations (transgenerational plasticity), allows organisms and their progeny to adapt to the environment without modification of the underlying DNA. Recent findings suggest that epigenetic modifications are important mediators of such plasticity. However, empirical studies have, so far, mainly focused on p...

Aim The species pool specific for a site includes all species from the region that are theoretically able to live in the site's particular ecological conditions. The absent portion of the site‐specific species pool forms the site’s dark diversity, which is unobservable and can only be estimated. Most existing methods to designate dark diversity ac...

Functional diversity assessments are crucial and increasingly used for understanding ecological processes and managing ecosystems. The functional diversity of a community is assessed by sampling traits at one or more scales (individuals, populations, species) and calculating a summary index of the variation in trait values. However, it remains uncl...

Heterostyly is a genetically determined floral polymorphism of style length promoting outcrossing between individuals of different morphs, which usually coexist within populations at equal frequencies. Loss in the area and connectivity of suitable habitats may cause deviations from the expected equal morph frequencies. However, there is a need to e...

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved....

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved....

Overyielding, the primary metric for assessing biodiversity effects on ecosystem functions, is often partitioned into “complementarity” and “selection” components, but this reveals nothing about the role of increased resource use, resource-use efficiency, or trait plasticity. We obtained multiple overyielding values by comparing productivity in a f...

Phenotypic plasticity is essential for organisms to adapt to local ecological conditions. Little is known about how mutualistic interactions, such as arbuscular mycorrhizal (AM) symbiosis, mediate plant phenotypic plasticity and to what extent this plasticity may be heritable (i.e. transgenerational effects). We tested for plant plasticity within-...

Agricultural intensification is one of the main drivers of species loss worldwide, but there is still a lack of information about its effect on functional diversity of arable weed communities. Using a large‐scale pan European study including 786 fields within 261 farms from eight countries, we analysed differences in the taxonomic and functional di...

Although one quarter of the species of plants and vertebrates are threatened with extinction, little is known about how the potential effect of extinctions on the global diversity of ecological strategies. Using trait and phylogenetic information for more than 75,000 species of plants, mammals, birds, reptiles, amphibians and freshwater fishes, we...

Functional redundancy is considered a major component of the insurance mechanism, which theoretically maintains ecosystem stability by preventing the loss of ecosystem functions with species loss. Over the past decades, examination of functional trait patterns to elucidate processes of community stability and ecosystem functioning have stimulated c...

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Understanding biodiversity changes in the Anthropocene (e.g. due to climate and land‐use change) is an urgent ecological issue. This important task is challenging because global change effects and species responses are dependent on the spatial scales considered. Furthermore, responses are often not immediate. However, both scale and time delay issu...

Saproxylic beetles depend on diverse microhabitats that are exploited by different species assemblages. We focused on anlyse the functional trait patterns and functional diversity components of two main assemblages that were collected with window traps (WTs) and hollow emergence traps (HETs) respectively, between three protected areas of the Iberia...

Functional diversity (FD) has the potential to address many ecological questions, from impacts of global change on biodiversity to ecological restoration. There are several methods estimating the different components of FD. However, most of these methods can only be computed at limited spatial scales and cannot account for intraspecific trait varia...

Both intra‐ and interspecific differences in traits may modulate interactions between plants. Two mechanisms are hypothesized to regulate these effects: competitive hierarchies and trait dissimilarities, but it is unclear how the prevalence of each might depend on environmental conditions and on intra and interspecific differences. We sowed six rep...

Functional and phylogenetic diversity (FD and PD respectively) of the resident community are expected to exert a key role in community resistance to colonization by surrounding species, and their establishment success. However, few studies have explored this topic experimentally or evaluated the interactive effects of these diversity measures. We i...

DarkDivNet is a global research collaboration which explores the dark diversity— the set of species that are absent from a site despite being suitable under the site conditions and present in the region. Participants of the network survey vascular plant diversity both at local (10x10 m) and regional scales (radius 10 km) using a standardized approa...

The biodiversity of a site includes the absent species from the region that are theoretically able to live in the site’s particular ecological conditions. These species constitute the dark diversity of the site. Unlike present species, dark diversity is unobservable and can only be estimated. Most existing methods to designate dark diversity act in...

Traditionally, trait‐based studies have explored single‐trait‐fitness relationships. However, this approximation in the study of fitness components is often too simplistic, given that fitness is determined by the interplay of multiple traits, which could even lead to multiple functional strategies with comparable fitness (i.e. alternative designs)....

Questions: How do the traits of pastoral species respond to growth in mixture, nitrogen addition and season? What are the impacts of trait plasticity on community aggregate trait values? Study site: A large-scale field experiment on intensively managed dairy pastures in New Zealand. Methods: We measured traits linked to rate of return on investment...

Despite their importance, how plant communities and soil microorganisms interact to determine the capacity of ecosystems to provide multiple functions simultaneously (multifunctionality) under climate change is poorly known. We conducted a common garden experiment using grassland species to evaluate how plant functional structure and soil microbial...

Plant invasions can drastically change the structure of native communities, but it is not fully understood whether alien species occupy phylogenetic and functional space within the range occupied by natives, or provide a novel set of evolutionary origins and traits to the invaded communities. Here, we evaluated this open question with data on a lar...

Experimental demethylation of plant DNA enables testing for epigenetic effects in a simple and straightforward way without the use of expensive and laborious DNA sequencing. Plants are commonly demethylated during their germination with the application of agents such as 5‐azacytidine (5‐azaC). However, this approach can cause unwanted effects such...

Despite its negative impacts on the environment and biodiversity, tree plantations can contribute to biodiversity conservation in fragmented landscapes, as they harbor many native species. In this study, we investigated the impact of Eucalyptus plantations on the taxonomic and functional diversity of ant communities, comparing ant communities sampl...

Tools to calculate trait probability density functions (TPD) at any scale (e.g. populations, species, communities). TPD functions are used to compute several indices of functional diversity, as well as its partition across scales. These indices constitute a unified framework that incorporates the underlying probabilistic nature of trait distributio...

Extensive livestock grazing is a global land use activity that has numerous social, cultural, and environmental benefits. Many marginal zones grazed for centuries are now suffering from abandonment, which has been found to be interrelated with species and functional diversity loss, and has potential consequences for the provision of ecosystem servi...

Functional traits and phylogeny offer different, and often complementary, information about ecological differences between species, an essential step to uncover biodiversity assembly mechanisms and their feedbacks to ecosystem functions. However, traits and phylogeny are often related due to underlying trait evolution. Consequently, when combined,...

Habitat partitioning is a common ecological mechanism to avoid competition among coexisting species, and the introduction of new species into existing assemblages can increase competitive pressures. However, situations of species in allopatry and sympatry only differing in species presence but not in environmental conditions are scarce. Thus, disce...