Atlantic Forest Research Articles

New discoveries for the genus Miconia (Melastomataceae): a new species for the brazilian Atlantic Forest

We describe Miconia leonorae, a new species exclusive to eastern Brazil, mainly from the evergreen forests of the Atlantic Forest Domain. This species can be distinguished from a wider circumscription of Miconia lepidota. It is classified as Miconia supersect. Discolores sect. Multispicatae and can be distinguished from similar species by features such as an acute leaf base and apex, light brown to gray abaxial leaf blade indumentum with lepidote-stellate trichomes (becoming glabrescent on mature leaves), cylindrical, scorpioid inflorescences with basal branches only bifid or trifid, and an androecium with isomorphic stamens.

Investigation of Rabies virus in wild mammals of the atlantic forest in Rio de Janeiro, Brazil.

With the successful control of rabies transmitted by dogs in Brazil, wild animals have played a relevant epidemiological role in the transmission of rabies virus (RABV). Bats, non-human primates and wild canines are the main wild animals that transmit RABV in the country. It is worth highlighting the possibility of synanthropic action of these species, when they become adapted to urban areas, causing infections in domestic animals and eventually in humans. This work aimed to evaluate the circulation of RABV in the Pedra Branca Forest, an Atlantic Forest area, located in the state of Rio de Janeiro, Southeast Brazil. Saliva and blood samples were obtained from 60 individuals of eight species of bats, captured with mist nets, and 13 individuals of callitrichid primates, captured with tomahawk traps. Saliva samples were subjected to Reverse Transcription Polymerase Chain Reaction (RT-PCR), targeting the RABV N gene, with all samples being negative. Blood samples of all animals were submitted to the Rapid Fluorescent Focus Inhibition Test (RFFIT) to detect neutralizing antibodies (Ab) for RABV. Six bat samples (8%) were seropositive for RABV with antibody titers greater than or equal to 0.1 IU/mL. The detection of Ab but not viral RNA indicates exposure rather than current RABV transmission in the analyzed populations. The results presented here reinforce the importance of serological studies in wildlife to access RABV circulation in a region.

Neomensularia meridionalis sp. nov. (Hymenochaetales, Basidiomycota), a New Species from South America

During field surveys in the Atlantic Forest of Southern Brazil, specimens of a poroid and pileate species of Hymenochaetaceae were found and are described as a new species, Neomensularia meridionalis. This species is morphologically and phylogenetically (ITS and nLSU) close to Neomensularia crocitincta, distributed in Central America, Cuba and Jamaica, but can be distinguished by its larger basidiomata, a duplex context featuring a distinct black line, and golden yellow basidiospores. Additionally, an updated identification key for Neomensularia species worldwide is provided.

Monitoring agricultural drought using different indices based on remote sensing data in the Brazilian biomes of Cerrado and Atlantic Forest.

Several remote sensing indices have been used to monitor droughts, mainly in semi-arid regions with limited coverage by meteorological stations. The objective of this study was to estimate and monitor agricultural drought conditions in the Jequitinhonha Valley region, located in the Brazilian biomes of the Cerrado and Atlantic Forest, from 2001 to 2021, using vegetation indices and the meteorological drought index from remote sensing data. Linear regression was applied to analyze drought trends and Pearson's correlation coefficient was applied to evaluate the relationship between vegetation indices and climatic conditions in agricultural areas using the Standardized Precipitation Index. The results revealed divergences in the occurrences of regional droughts, predominantly covering mild to moderate drought conditions. Analysis spatial of drought trends revealed a decreasing pattern, indicating an increase in drought in the Middle and Low Jequitinhonha sub-regions. On the other hand, a reduction in drought was observed in the High Jequitinhonha region. Notably, the Vegetation Condition Index demonstrated the most robust correlation with the Standardized Precipitation Index, with R values ​​greater than 0.5 in all subregions of the study area. This index showed a strong association with precipitation, proving its suitability for monitoring agricultural drought in heterogeneous areas and with different climatic attributes. The use of remote sensing technology made it possible to detect regional variations in the spatio-temporal patterns of drought in the Jequitinhonha Valley. This vision helps in the implementation of personalized strategies and public policies, taking into account the particularities of each area, in order to mitigate the negative impacts of drought on agricultural activities in the region.

Antibodies against Leptospira spp. in Free-living and Captive Broad-snouted Caiman (Caiman latirostris) and Free-living Yacare Caiman (Caiman yacare) in Brazil.

We evaluated antibodies against Leptospira spp. in both free-living and captive Caiman latirostris from Atlantic Forest, and free-living Caiman yacare from Pantanal, Brazil, by using a microscopic agglutination test. Overall seropositivity was 17%, with rates of 36% in captive C. latirostris (n=4/11) and 18% in free-living C. yacare (n=4/22).

Land cover effects on regenerants density and richness in restoration treatments in the Atlantic Rainforest biome

There is good evidence that the results of a restoration program depend largely on the landscape context. In restoration projects, it is crucial to consider the previous land uses and covers and landscape configuration as they have a significant impact on the entire process, especially seed dispersal and natural regeneration of sites. Thus, the objective of this work was to verify the effect of landscape (i.e., land use) on restoration success of total planting, seed sowing and natural regeneration sites in the Atlantic Forest biome, southeast Brazil. The methodology employed was based on demonstration units of restoration. The focus was on regenerant richness and abundance indicators, along with land use and land cover data (class area) from 2010, 2015, and 2020, obtained from the MapBiomas Project. Generalized linear models and correlation analysis were used for the study. We hypothesize that a landscape with more forest and natural regeneration cover (mosaic of agriculture and pasture) would positively affect regenerants in restored sites, which we only observed for natural regeneration treatment sites. In active restoration treatment sites, mosaic of agriculture and pasture was negatively and farming was positively associated to regenerants. We also found greater density and richness of regenerants in natural regeneration treatment sites than in seed sowing and seedling planting. The influence of the surrounding landscape, particularly mosaic, played a crucial role in this success. In addition, land use cover history, as we observed in classes in the last five and ten years, did explain regenerants in studied sites. Thus, results show that the characterization of landscape context and previous land-use history is essential to understand the limitations to succession and define cost-effective restoration strategies.

Unlocking the potential of uvaia (Eugenia pyriformis Cambess): exploring its rich chemical composition and varied biological activities for innovations in the food industry

SummaryEugenia pyriformis Cambess (uvaia), a native species of the Atlantic Forest ecosystem, has received limited attention in scientific investigations. Renowned for its distinctive organoleptic characteristics, this fruit is suitable for immediate consumption and industrial processing. Additionally, it is rich in vitamins and bioactive compounds, which offer potential health benefits. These bioactive compounds, such as ascorbic acid, carotenoids, and phenolic compounds, have demonstrated antioxidant and antimicrobial properties in both in vitro and in vivo studies. This review consolidates recent research on uvaia, focusing on its chemical composition, biological activities, and potential applications within the food industry. It aims to lay the groundwork for future research and promote its recognition and utilisation in food production.

The role of artificial ponds for anuran diversity in managed areas of the Atlantic Forest

ABSTRACT Artificial ponds can be important reproductive sites for anurans in human-disturbed habitats. However, this capacity depends on the configuration of the habitat where they are inserted. We evaluated how anuran beta diversity was influenced by the configuration of the surrounding habitat and water parameters in artificial ponds in southern Brazil. Physicochemical parameters of water and the configuration of the surrounding habitat were the elements that best explained the variation in beta diversity. However, the surrounding habitat had a predominant role in the differences between communities. The level of exposed soil was the component that best acted on the turnover component of beta diversity followed by urbanization, which highlights the impact of human presence on anuran diversity. In other words, we observed that anthropogenic changes caused changes in species composition in the evaluated ponds. Among water parameters, only phosphate was relevant for the configuration of communities. Phosphate peaks may result from sewage or fertilizer contamination, not infrequent in the region despite the low human density and the small-scale agriculture economy. Even though they are anthropogenic environments, artificial ponds can help maintain the anuran communities, but their effectiveness will depend on the presence of forest remnants and water quality.

Ecological inferences in Orchidaceae species from the Brazilian subtropical Atlantic Forest based on morphological and functional anatomical traits

Ecological anatomy studies seek to link structural aspects to environmental pressures, improving knowledge about the occurrence and distribution of species. Orchidaceae is a group with high species richness and ecological diversity, well represented in the Brazilian Atlantic Forest, a biodiversity hotspot. The Iguaçu National Park (ParNa Iguaçu) is an important remnant of the southern Brazilian Atlantic Forest with a high richness of Orchidaceae. Given the family's diversity and the relevance of ecological anatomy studies for interpreting traits associated with the survival of plants in certain environments, we studied 38 species of Orchidaceae from ParNa Iguaçu, seeking to identify possible functional groups. For this, we surveyed 57 leaf morpho-anatomical characters and conducted histochemical and statistical tests. The morpho-anatomical traits group the species into two functional groups, with Cyclopogon congestus emerging isolated. In group A, xeromorphic morphoanatomical characters predominate, and group B combines members with meso/hygromorphic characteristics. We advocate the importance of understanding the ecological behavior of species, especially Orchidaceae, due to their important ecological role, in conservation actions in protected areas. Furthermore, our results highlight the morphological and anatomical diversity of the Orchidaceae found in ParNa Iguaçu, as well contributing to the development of better management practices for these plants, considering the threats posed by the opening of clearings and climate change.

Impacts of urbanization on multiple dimensions of bird diversity in Atlantic forest landscapes

Urbanization is resulting in habitat loss and fragmentation around the world, but its effects on biodiversity remain poorly understood. We investigated how the increase of urban cover surrounding nine Brazilian Atlantic forest reserves (14 ha - 1058 ha) affects the taxonomic, functional and phylogenetic diversity of bird communities, considering forest and non-forest species separately. Using a multi-scale local landscape approach, we estimated the percentage of urban cover in circular buffers of varying radii (200–1000 m). We calculated 10 complementary metrics of bird diversity and modelled them against the percentage of urban cover using generalized additive models. Urbanization had a negative, non-linear impact on the taxonomic diversity of bird communities, with a sharp decrease of up to 50 % in the effective number of rare, typical and dominant species in landscapes with 20–40 % of urbanization. This decrease was not restricted to forest species (urban avoiders), but also affected non-forest ones (urban utilizers and dwellers). The impact of urbanization on functional and phylogenetic diversity were less evident, suggesting that the vulnerability to urbanization has a weak phylogenetic and phenotypic signal. We conclude that urbanization impacts a broad spectrum of bird clades and traits and drastically reduce the taxonomic diversity of bird communities.

New species of the long-horned caddisfly Oecetis McLachlan, 1877 (Trichoptera: Leptoceridae) from the Atlantic Forest, Brazil and their evolutionary relationship

Asymmetrical genitalia are reported from majorTrichoptera subgroups and evolved multiple times independently. In Oecetis, it is a characteristic of the insconpicua group. However, certain species in other species groups also evolved an asymmetrical spiny projection on the phallotheca. Here, two new species with an asymmetric projection are described in the falicia group from the Brazilian Atlantic Forest, Espírito Santo state. Their phylogenetic relationships were investigated through a Bayesian analysis combining COI and morphological data. Additionally, we provide new records of Oecetis connata, O. inconspicua and O. paranensis from the Espírito Santo state, and O. connata and O. flinti from Minas Gerais state. Oecetis capixaba sp. nov. is placed with low support as the sister species of O. acanthostema, both presenting stout spine-like setae on the inner surface of the inferior appendage; the new species is diagnosed by the long spine-like setae on the inferior appendage, the narrow dorsolateral process of segment IX, and the phallic apparatus without apical projections. Oecetis ruschii sp. nov. is placed as a sister group of the clade including O. facilia and O. furcata, both presenting forked dorsolateral processes of segment IX; the new species is diagnosed by the dorsolateral process of segment IX with a lateral branching and the apex of inferior appendage wide and triangular. While the phylogenetic results should be considered preliminary and interpreted with caution, they indicate that the asymmetric projection evolved multiple times convergently in the avara, punctata, and falicia groups. The asymmetric genitalia in the falicia group seems to have evolved only in males since no correspondent asymmetry is described for females. The function of the asymmetrical projection remains unknown.

An evolutionary disruption of the buzz pollination syndrome in neotropical montane plants.

Under pollinator limitations, specialized pollination syndromes may evolve toward contrasting responses: a generalized syndrome with increased pollinator attraction, pollinator reward, and pollen transfer capacity; or the selfing syndrome with increased self-pollen deposition, but reduced pollinator attraction and pollen transfer capacity. The buzz-pollination syndrome is specialized to explore female vibrating bees as pollinators. However, vibrating bees become less-active pollinators at montane areas of the Atlantic Forest (AF) domain. This study investigated whether the specialized buzz-pollination syndrome would evolve toward an alternative floral syndrome in montane areas of the AF domain, considering a generalized and the selfing syndromes as alternative responses. We utilized a lineage within the buzz-pollinated Miconia as study system, contrasting floral traits between montane AF-endemic and non-endemic species. We measured and validated floral traits that were proxies for pollinator attraction, reward access, pollen transfer capacity, and self-pollen deposition. We inferred the evolution of floral trait via phylogenetic comparative methods. AF-endemic species have selectively evolved greater reward access and more frequently had generalist pollination. Nonetheless, AF-endemic species also have selectively evolved toward lower pollen transfer capacity and greater self pollination. These patterns indicated a complex evolutionary process that has jointly favored a generalized and the selfing syndromes. The buzz pollination syndrome can undergo an evolutionary disruption in montane areas of the AF domain. This floral syndrome is likely more labile than often assumed, allowing buzz-pollinated plants to reproduce in environments where vibrating bees are less-reliable pollinators.

Hidden diversity of Aegla (Decapoda, Anomura) in a priority area for conservation

AbstractAegla is a genus of freshwater anomuran crabs encompassing 94 described species occurring in meridional South America. Brazil harbours the greatest diversity of species, mainly in its southern region, which becomes a priority area for Aegla conservation due to high species richness and phylogenetic diversity, elevated endemism and threats to the Atlantic Forest. Here, we assessed Aegla richness and distribution in this priority area and tested the hypothesis of crypticism in the region's species. Sampling sites were distributed inside and in the surroundings of five protected areas, covering the major distribution range of several species described for southern Brazil. We applied three species delimitation methods to COI mitochondrial gene data, coupled with phylogenetic and morphological analyses. By integrating species delimitation based on the COI data and traditional taxonomy, we uncovered a high proportion of hidden diversity among Aegla in our study area and identified ten known species besides impressive 17 new putative species. Of these, eight are possible new taxonomic units that do not conform to any known species, and nine are divided into three cryptic groups: Aegla jarai complex, Aegla franciscana complex and Aegla camargoi complex. Most of these species have a narrow spatial distribution, making them highly vulnerable to habitat degradation. Despite the real richness of the genus still being largely unknown, the high diversity revealed here makes our study area the richest known region regarding the number of Aegla species within its distribution, making this area even more worthy for conservation efforts.

Spatial bias in sampling small rodents in the Atlantic Forest: A landscape and accessibility perspective

Understanding the effects of habitat loss and fragmentation on species spatial distribution is challenging, mainly because knowledge of species occurrences is biased. Survey efforts are unevenly distributed causing spatial sampling biases that are normally neglected. Assessing sampling bias is particularly urgent for threatened ecoregions, such as the Atlantic Forest, a global biodiversity hotspot. Here, we assess spatial sampling biases of small rodents in the Atlantic Forest, using an integrative approach with accessibility and landscape metrics. We built a robust dataset of 11,495 primary records of the Atlantic Forest’s small rodent species, based on information from digitally accessible repositories. We expect that well-sampled sites are spatially aggregated and nearer roads, urban centers, on landscapes with larger forest fragments, and with higher percentage of forest cover. We also expect gaps of small rodents sampling in rare landscape conditions. Our results indicated that only less than 1% of the Atlantic Forest (at 1 km2 cell-size resolution) are well sampled. Following our expectations, the well-sampled sites were spatially aggregated biased toward roads, urban centers, larger forest fragments, and landscapes with higher percentage of forest cover. We also found a survey gap on common landscape conditions. Our findings suggest that the spatial distribution of small rodents at landscape level (1 km2) remains unknown across most of the Atlantic Forest spatial extension. Our findings also point to new priority sites for small mammals sampling on common landscape conditions, in smaller fragments and on remote areas improving spatial distribution knowledge and contributing to conservation policies at landscape level.

Elevational shifts in tree community composition in the Brazilian Atlantic Forest related to climate change

AbstractQuestionClimate change induces shifts in species distributions, ultimately changing community composition. Mountains are especially sensitive to climate change, and tree species are predicted to move towards higher elevations, but observed changes are not always unidirectional. The diversity and complexity of tropical and subtropical systems limits our understanding of climate‐induced responses of subtropical mountain forests. Here, we investigated migration trends in tree community composition along an elevational gradient, and between the transition from lowland to montane forests in subtropical forests.LocationBrazilian Atlantic Forest.MethodsWe used thermal affiliations of 627 tree species to calculate community temperature scores (CTS) for different life‐history stages of trees in 96 permanent plots. We compared CTS of different life‐history stages across space and time.ResultsMost tree communities (58%) did not show a significant difference of CTS between life‐history stages, indicating a non‐migration trend. On the other hand, 27% of tree communities showed upward migration and 15% downward migration. Upward migration was more common in montane forests, and downward migration in lowland forests. Our temporal analysis shows significant changes in CTS values for juvenile communities with 0.36°C decrease in lowland forests and 0.34°C increase in montane forests.ConclusionsContrasting results between lowland and montane forest communities indicate that the transition zone influences migration patterns and may reflect differences in species’ thermal limitations, as well as by non‐thermal factors such as biotic interactions. Our findings provide the first evidence of climate change‐induced community shifts in the Brazilian Atlantic Forest. We demonstrated that upward migration trends were predominantly observed in montane–upper mountain forests, while downward migrations were noted in lowland–submontane forests. The thermophilization of montane forests may suggest an increased risk of reduction for cold‐demanding species under climate change scenarios.

Brocket deer niche breadth and overlap: spatial similarities limit species coexistence

AbstractUnderstanding species coexistence requires information about how species interact and partition their ecological niche. Niches of closely related species tend to be quite similar and thereby, when syntopic, overlap can result in competition when shared resources are limiting. Brocket deer of tropical forests are ecologically similar species due to convergence or relatedness. As a group, what little is known about their ecology suggests spatial and temporal niche separation while dietary differences among species remain largely unknown. The possibility of competition has been suggested, as it is rare for more than two species to co‐occur. If competition is important, then it may limit syntopy of more species and its importance should increase with relatedness between species. Based on analyses of faecal samples, camera traps, and stable isotopes and multidimensional analysis of niche space, we test spatial, temporal and dietary niche differences and compare niche breadth in two syntopic species pairs: Mazama rufa with M. nana (both of subtribe Odocoileina) and M. jucunda (Odocoileina) with Subulo gouazoubira (Blastocerina). In 2007–2008 and 2017–2019, 240 faecal samples and 375 records were collected in the Brazilian Atlantic Forest. Spatial partitioning among both species pairs was found, with S. gouazoubira and M. nana having greater niche breadth, which may permit or facilitate coexistence. Odocoileina tend to be nocturnal, while Blastocerina tend to be diurnal, which suggests that phylogeny may influence species coexistence. Furthermore, the species tended to overlap with respect to diet. These results suggest interference competition avoidance by niche partitioning at the level of habitat and time, allowing coexistence. Thereby, despite the morphological similarities, brocket deer species of evolutionarily convergent subtribes preserve differences in ecological niche dimensions that facilitate the coexistence of their populations.

Larval morphology, vocalisations, and phylogenetic position of Crossodactylus dantei (Anura: Hylodidae): new insights into a little-known anuran species

Aspects of the biology and natural history of the endemic anuran Crossodactylus dantei remain unknown even almost 30 years since its description. In this study, we describe its larval stage, along with the advertisement and territorial calls, and assess its phylogenetic positioning based on a fragment of the 16S rRNA mitochondrial gene. The larval stage of C. dantei follows the morphological pattern of Hylodidae, corroborating the relatively uniform larval morphology of the family. However, it is still possible to observe characters that distinguish C. dantei from other species in the genus, such as the elliptical body in dorsal view, nostrils located closer to the snout than to the eyes, an elongated pre-nasal arena, and the absence of taenia tecti medialis and taenia tecti transversalis in the chondrocranium. Its advertisement call is characterised by a sequence of about 23 pulsed notes, with ascending amplitude modulation, and has a mean duration of 2.5 seconds, with a dominant frequency of 3356 Hz. The territorial call has a mean duration of 23 seconds, a dominant frequency of 3160 Hz, and two distinct types of notes. Phylogenetic analysis indicated the existence of two main lineages in the genus. Crossodactylus dantei was recovered with high statistical support as a sister species to the clade composed of C. gaudichaudii and C. timbuhy. Furthermore, the group formed by these three species was recovered as the sister lineage of C. trachystomus. In summary, the information presented here represents a significant advance in the knowledge of this enigmatic and rare species of the northern Atlantic Forest, providing baseline data for future ecological and evolutionary studies of this unique frog. Keywords: tadpole, oral anatomy, chondrocranium, calls, phylogeny, Atlantic Forest

Overcoming biases of birds research in the Caatinga

“The biodiversity of the Caatinga is poorly known” has been a mantra among studies on biodiversity in Brazil, including birds. However, species richness in semiarid regions is expected to be lower than that in other ecosystems, and studies in the Caatinga have increased in recent decades. Therefore, this narrative may not only reflect knowledge bias. Here, we gathered complementary datasets of bird assemblages from literature and an online database of the Caatinga and other phytogeographic domains in Brazil. We then employed novel spatial and temporal predictive statistics to address this question. We estimate that the Caatinga is only six species short of fully knowing its taxonomic diversity. We did find important spatial knowledge gaps regarding species distribution, but only higher than that of the Atlantic Forest. The species richness and distribution of Caatinga birds are as well-known as the Pampa and Pantanal and are better known than the Cerrado and the Amazon rainforest. Our findings challenge the notion that the Caatinga is the poorest known region regarding birds. Bird research in the region should focus on advancing a research and conservation agenda that enhances understanding of regional biodiversity and ecosystem dynamics while promoting ecological syntheses for tropical dry forests.

Montane diversification as a mechanism of speciation in neotropical butterflies.

The mountains in the Atlantic Forest domain are environments that harbor a high biodiversity, including species adapted to colder climates that were probably influenced by the climatic variations of the Pleistocene. To understand the phylogeographic pattern and assess the taxonomic boundaries between two sister montane species, a genomic study of the butterflies Actinote mantiqueira and A. alalia (Nymphalidae: Acraeini) was conducted. Analyses based on partial sequences of the mitochondrial gene COI (barcode region) failed to recover any phylogenetic or genetic structure discriminating the two species or sampling localities. However, single nucleotide polymorphisms gathered using Genotyping-by-Sequencing provided a strong isolation pattern in all analyses (genetic distance, phylogenetic hypothesis, clustering analyses, and F ST statistics) which is consistent with morphology, separating all individuals of A. alalia from all populations of A. mantiqueira. The three sampled mountain ranges where A. mantiqueira populations occur-Serra do Mar, Serra da Mantiqueira, and Poços de Caldas Plateau-were identified as three isolated clusters. Paleoclimate simulations indicate that both species' distributions changed according to climatic oscillations in the Pleistocene period, with the two species potentially occurring in areas of lower altitude during glacial periods when compared to the interglacial periods (as the present). Besides, a potential path between their distribution through the Serra do Mar Mountain range was inferred. Therefore, the Pleistocene climatic fluctuation had a significant impact on the speciation process between A. alalia and A. mantiqueira, which was brought on by isolation at different mountain summits during interglacial periods, as shown by the modeled historical distribution and the observed genetic structure.

Detection of functional diversity gradients and their geoclimatic filters is sensitive to data types (occurrence vs. abundance) and spatial scales (sites vs. regions)

Functional diversity (FD) reflects within- and between-site variation of species traits (α- and β-FD, respectively). Understanding how much data types (occurrence-based vs. abundance-weighted) and spatial scales (sites vs. regions) change FD and ultimately interfere with the detection of underlying geoclimatic filters is still debated. To contribute to this debate, we explored the occurrence of 1690 species in 690 sites, abundances of 1198 species in 343 sites, and seven functional traits of the Atlantic Forest woody flora in South America. All FD indices were sensitive and dependent on the data type at both scales, with occurrence particularly increasing α richness and dispersion (occurrence > abundance in 80% of the sites) while abundance increased β total, β replacement, and α evenness (abundance > occurrence in 60% of the sites). Furthermore, detecting the effect of geoclimatic filters depended on the data type and was scale-dependent. At the site scale, precipitation seasonality and soil depth had weak effects on α- and β-FD (max. R2 = 0.11). However, regional-scale patterns of α richness, dispersion, and evenness strongly mirrored the variation in precipitation seasonality, soil depth, forest stability over the last 120 kyr, and cation exchange capacity (correlations > 0.80), suggesting that geoclimatic filters manifest stronger effects at the regional scale. Also, the role of edaphic gradients expands the idea of biogeographical filters beyond climate. Our findings caution functional biogeographic studies to consider the effect of data type and spatial scale before designing and reaching ecological conclusions about the complex nature of FD.