Fern and Lycophyte Evolution Lab

Federal University of Pernambuco | Evolutionary Biology | Ferns and Lycophytes

Our Publications

2026

A comparison of interspecific and intraspecific phenotypic variation in spectral signatures of ferns with robust versus uncertain species boundaries

Authors: Niksoney A. Mendonça | Marise H. V. Oliveira | Thais E. Almeida

Journal: Journal of Plant Research (2026) - Original research article

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Abstract: Species delimitation methods based on macromorphology are often limited by phenotypic plasticity in plants. FourierTransform near-infrared spectroscopy (FT-NIR) provides a promising alternative as a non-destructive technique that mea-sures molecular vibrations (overtone and combination bands of C–H, N–H, and O–H bonds) from plant tissue exposed tonear-infrared light (780–2,500 nm). We applied FT-NIR to the taxonomically challenging Scaly clade of Microgramma ferns (94 samples, eight species), including dimorphic and monomorphic taxa, to evaluate its diagnostic potential. Usingmultivariate models and cross-validation, we achieved 81–100% average identification accuracy. Well-defined species(e.g., M. percussa) reached 100% accuracy, while morphologically overlapping taxa showed lower accuracy, likely due tohybridization, introgression, or cryptic variation. Dimorphic species exhibited higher intraspecific spectral variability andlower accuracy linked to differences between fertile/sterile fronds than monomorphic species. FT-NIR proves effective asa complementary tool for fern systematics, elucidating species limits and diversity patterns. Further studies should addresshow hybridization, introgression, and indumentum affect spectral data.

Ferns FT-NIR spectroscopy Microgramma Predictive models Species limits Systematic
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Mesoamerican roots: the biogeographic history of the fern genus Pleopeltis

Authors: Carolina do Valle | Alexandre Salino | Lucas V. Lima | Ingridy O. Moura | Thais E. Almeida

Journal: Botanical Journal of the Linnean Society (2026) - Original research article

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Abstract: The genus Pleopeltis stands out for its high species richness in tropical montane regions and its disjunct neotropical–afrotropical distribution, with most species occurring in the Neotropics. This distribution provides a unique opportunity to explore dispersal patterns, a key mechanism in fern biogeography. In this study, we aimed to identify the biogeographic processes shaping the distribution of Pleopeltis and the directionality of its dispersal events throughout evolutionary history. We analysed a combined dataset of three plastid regions from 70 species to infer divergence times. Ancestral range estimation and stochastic biogeographic mapping were used to elucidate biogeographic patterns and processes. Our findings showed that dispersal was the main process driving the genus’s distribution, accounting for nearly half of the inferred biogeographic events. Notably, we identified a recent west-to-east dispersal pattern connecting the Neotropics with the Paleotropics, highlighting the significance of long-distance dispersal in ferns. Mexico and Central America were identified as the centre of origin and diversity for the genus and the main source of dispersal. The Andes were the second most important centre, functioning as a source and sink for dispersal events. These findings emphasize the critical roles of these dynamic centres in the diversification and distribution of global biodiversity.

dispersal directionality long-distance dispersal phylogenetic inferences source-sink
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2025

A new generic name to solve the paraphyly of Sticherus C.Presl (Gleicheniaceae, Polypodiopsida)

Authors: Lucas V. Lima | Alexandre Salino | Germinal Rouhan | Thaís E. Almeida

Journal: Phytotaxa (2025) - Original Research Article

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Abstract: Gleicheniaceae is an early diverging lineage of leptosporangiate ferns, characterized by pseudodichotomous frond branching. While six genera are phylogenetically supported, Sticherus has been found to be paraphyletic. To resolve this issue, we describe a new genus, Sticheroides gen. nov., to accommodate Sticherus truncatus and S. milnei, based on phylogenetic placement, time divergence depth, and frond architecture. We provide an updated identification key to the genera, descriptions, and comments. Sticheroides diverged from the clade formed by Gleichenia+Stromatopteris+Sticherus approximately 114–117 Mya and is distinguished by buds and rhizomes covered with scales and anisotomic branching. The new genus solves the paraphyly of Sticherus as defined so far, while ensuring nomenclatural stability within Gleicheniaceae.

Gleicheniales Leptosporangiate ferns Nomenclature New taxon Taxonomy
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Blowin' in the Wind: Ferns and Lycophytes Connect Biogeographic Refuges in the Atlantic Forest

Authors: D. Victor S. e Silva | Lucas V. Lima | Mércia P. P. Silva | Thais E. Almeida | Augusto C. P. Santiago

Journal: Journal of Biogeography (2025) - Original Research Article

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Abstract: Test the floristic structure of fern and lycophyte in the Northeastern Atlantic Forest based on the two sub-regions of endemism, (1) montane inland wet forests and (2) Coastal Forests. Northeastern Brazil's Atlantic Forest, particularly the Coastal and Montane inland wet forests. Floristic composition was compiled from GBIF data, and environmental data were extracted from rasters for 57 forest remnants. PCA analyses were carried out to assess environmental structuring, NMDS to test floristic dissimilarity, and generalised dissimilarity modelling (GDM) to examine environmental effects on community distribution. Clusters were formed by K-means, and statistical analyses such as PERMANOVA and ANOSIM were carried out to test the significance of the results. A total of 307 species were recorded, 93.2% of which were ferns and 6.8% lycophytes. The floristic composition showed 71.6% of species shared between both sub-regions. PCA displayed significant environmental differences between montane inland wet forests and coastal forests, but NMDS did not indicate any separation in floristic composition. However, the clusters identified by K-means were highly significant (R = 0.445), in contrast to the groupings of the sub-regions (R = 0.124). GDM explained 78.8% of the distribution pattern, and its cluster classification revealed no clear structure between NAF sub-regions. Additionally, GDM highlighted variables such as geographic distance, cloud cover, rainfall, silt content and soil pH as determinants of the communities. Despite differences in environmental factors between montane inland wet forests and coastal forests, the floristic composition of ferns and lycophytes does not reflect this separation, suggesting that historical biogeography and long-distance dispersal facilitate high connectivity between these sub-regions.

Biogeography Ferns Lycophytes Atlantic Forest Dispersal
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Rediscovery after three decades of the freshwater sponge Metania kiliani on a terrestrial fern

Authors: Carolina do Valle | Keven dos S. Lima | Ulisses Pinheiro | Thaís E. Almeida

Journal: Acta Amazonica (2025) - Short Communication

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Abstract: This study presents the second record of Metania kiliani (Porifera: Metaniidae) three decades after its description in 1992. The sponge was found on a herbarium specimen of the terrestrial fern Actinostachys pennula (Schizaeaceae), which is typically known to be absent from flooded environments. As sponges need at least some weeks to establish themselves on a support, this first record of a freshwater sponge on this fern suggests that A. pennula can withstand prolonged submergence. This is the first record of M. kiliani for the state of Roraima, Brazil.

Actinostachys Amazonia Metaniidae Porifera Schizaeaceae white-sand vegetation
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Ferns and lycophytes in the Cerrado: a floristic study in eastern Maranhão, northeastern Brazil

Authors: Niksoney A. Mendonça | Sirlane S. Oliveira | Juliana Aljahara | Thaís E. Almeida | Roziajne S. Fernandes

Journal: Rodriguesia (2025) - Original Research Article

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Abstract: The present study contributes to the knowledge of the fern and lycophyte flora of the city of Chapadinha, an area of the Cerrado located in the eastern state of Maranhão, Brazil. The study also presents taxonomic and environmental data on the occurrence of these species. Species were collected during five expeditions performed between December 2021 and August 2022, in two protected areas and some sites associated with wet environments outside these areas. In addition, specimens deposited in an herbarium were examined. In total, 20 species of ferns and three species of lycophytes were recorded, distributed over 12 families and 16 genera. The most representative families were Pteridaceae (six species) and Thelypteridaceae (four species). Despite the anthropization, riparian forests showed the highest richness (21 species), among which the non-flooded riparian forests stood out (18 species). Our data highlight the importance of identifying fern and lycophyte species and emphasize the potential of these species in future conservation studies and as ecological bioindicators.

bioindicators flora inventory vascular plants taxonomy
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2024

Isolated in biogeographic islands: discovery of a disjunct population of a fern in Northeastern Atlantic Forest

Authors: D. Victor S. e Silva | Niksoney A. Mendonça | Maria F. Barros | Thais E. Almeida | Augusto C. P. Santiago

Journal: Revista Peruana de Biología (2024) - Scientific note

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Abstract: Brejos Nordestinos (northeastern highland wet forests) is a unique ecosystem within the Northeastern Atlantic Forest. This area encompasses remmants of humid forests above 600 meters in the semi-arid region. These environmental conditions create an optimal ecosystem for the diversification of plants, including ferns. Pico do Jabre in Paraíba state, with an elevation of 1,208 meters above sea level, is an example of Brejo Nordestino. This study reports the occurrence of Pleopeltis pleopeltidis in the Northeastern Atlantic Forest, particularly at Pico do Jabre, a new record of P. pleopeltifolia for Paraíba state, and the local fern flora. Additional records of P. pleopeltidis were found in the municipalities of Triunfo and Floresta, in Pernambuco state, which are also classified as Brejos Nordestinos with elevations exceeding 1,000 m a.s.l. These findings extend the species known extent of occurrence by approximate 40%. The conservation status of P. pleopeltidis is suggested as Vulnerable (VU) across its entire range and Endangered (EN) in Northeast Brazil. The distance between populations in the Northeastern Atlantic Forest and the nearest populations in Bahia state ranges from 763–931 km, indicating a potential disjunction. Although the species occurs in three relatively close Brejos Nordestinos, the fragmented nature of the Northeastern Atlantic Forest results in isolated populations on these biogeo- graphical islands, surrounded by a Caatinga (seasonally dry tropical forest) matrix.

Brejos de Altitude Pico do Jabre Pleopeltis pleopeltidis Species conservation Species distribution
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So close no matter how far: The floristic affinities of the Serra do Papagaio ferns and lycophytes within the Atlantic Forest

Authors: Lucas V. Lima | Alexandre Lima | Vinícius A. O. Dittrich | Samyra G. Furtado | Luiz M. Neto | Thaís E. Almeida

Journal: Acta Botanica Brasilica (2024) - Original Research Article

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Abstract: We discuss the importance of fern and lycophyte species in the Atlantic Forest ecosystem as well as the need for floristic inventories to understand their distribution and conservation status. We focus on the Serra do Papagaio region in southeastern Brazil, which is known for its high plant diversity but lacks comprehensive inventories of ferns and lycophytes. In this study, we conducted a floristic inventory of Serra do Papagaio and identified 172 fern and 29 lycophyte species. Our findings reveal 21 threatened species and two new records for Minas Gerais State. Additionally, we compared the floristic composition of Serra do Papagaio with that of other areas within the Atlantic Forest using non-metric multidimensional scaling (NMDS) and generalized dissimilarity modeling (GDM) analyses. The results highlight distinct clusters of areas based on elevation and geographic location. We emphasize the need for further sampling and conservation efforts in Serra do Papagaio as well as provide valuable insights into the drivers of fern and lycophyte diversity in the Atlantic Forest.

Floristic survey Hotspot Serra da Mantiqueira Southeastern Brazil mountains
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The classification of the fern family Gleicheniaceae, with the description of a new genus, segregated from Sticherus

Authors: Lucas V. Lima | Thaís E. Almeida | Michael Kessler | Germinal Rouhan | Shuichiro Tagane | Vinícius A. O. Dittrich | Alexandre Salino

Journal: Botanical Journal of the Linnean Society (2024) - Original Research Article

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Abstract: Gleicheniaceae is an early diverging lineage of leptosporangiate ferns. Although the family can be readily recognized by the pseudodichotomousbranching pattern of its fronds, generic circumscription has long been debated. Phylogenetic analyses based on genomic data supported themonophyly of six genera (Dicranopteris, Diplopterygium, Gleichenella, Gleichenia, Rouxopteris, and Stromatopteris) but indicate the paraphylyof Sticherus. To accommodate this latter result, we describe a new genus, Sticheropsis, gen. nov., segregated from Sticherus. We provide a newtaxonomic treatment of Gleicheniaceae, including an identification key to all eight genera, descriptions of the genera with comments and notesconcerning geographic distributions and phylogenetic affinities, as well as an overview of the morphology of the family with a revised and uni-fied terminology for the fronds. We recognize 149 species plus two hybrids for the family, distributed into eight genera, and propose six newcombinations.

Gleicheniales Morphology New combinations Synopsis Taxonomy
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Editorial

Editorial: Biology, systematics, and evolution of ferns and lycophytes in the omics era, volume II

Authors: Li-Y. Kuo, Alexandre Salino & Thaís E. Almeida - Frontiers in Plant Science (2025)Access editorial

Editorial: The future of the homology concept under novel evolutionary paradigms

Authors: Thaís E. Almeida, Thiago André & Roxana Yockteng - Frontiers in Plant Science (2024)Access editorial

Nomenclature communications

(3006) Proposal to conserve the name Sticherus (Gleicheniaceae) with a conserved type

Authors: Lucas V. Lima, Thaís E. Almeida, Michael Kessler, Germinal Rouhan & Alexandre Salino - TAXON (2024)Access nomenclature comunications

Commentary

Recurrent allopolyploidy and its implications for conservation in vascular plants: a commentary on ‘Population genomics of the Isoetes appalachiana (Isoetaceae) complex supports a “diploids-first” approach to conservation’

Authors: Thaís E. Almeida & Bárbara S. S. Leal - Annals of Botany (2024)Access commentary

Dataset

Biodiversidade de samambaias e licófitas da Paraíba, Brasil

Authors: Sergio R. S. Xavier, D. Victor S. Silva, Thaís E. Almeida, Gabriela de L. D. Cavalcante, Antonio J. N. Silva, Hanna S. S. Neves, João V. M. Moreira & Augusto C. P. Santiago - DATAPB (2025)Access dataset

Preprint

Paths and obstacles to an evidence-based, reproducible, and dynamic Plant Taxonomy

Authors: Thiago Andre & Thaís E. Almeida — SciELO Preprints (preprint), 2025. Access preprint

Rediscovery after three decades of the freshwater sponge Metania kiliani on a terrestrial fern

Authors: Carolina do Valle M. de S., Keven dos S. Lima, Ulisses Pinheiro & Thaís E. Almeida — SciELO Preprints (preprint), 2025. Access preprint

Mesoamerican roots: the biogeographic history of the fern genus Pleopeltis

Authors: Carolina do Valle, Alexandre Salino, Ingridy O. Moura, Lucas V. Lima & Thaís E. Almeida — SciELO Preprints (preprint), 2025. Access preprint

Filling Gaps: Floristic Survey of Ferns and Lycophytes in Rio Grande do Norte state, northeastern Brazil

Authors: Maurício B. Nascimento, Augusto C. O. Santiago, Fernanda A. Carvalho & Thaís E. Almeida — SciELO Preprints (preprint), 2025. Access preprint

Blowin' in the wind: ferns and lycophytes connect biogeographic refuges in the Atlantic Forest

Authors: D. Victor S. Silva, Lucas V. Lima, Mércia P. P. Silva, Thaís E. Almeida & Augusto C. P. Santiago — SciELO Preprints (preprint), 2025. Access preprint

Fantastic plants and where to find them: niche occupancyanalysis in neotropical epiphytic species

Authors: Keven dos S. Lima, Lucas v. Lima & Thaís E. Almeida — SciELO Preprints (preprint), 2025. Access preprint

Spectral signatures as evidence to test hypotheses in plant species complexes

Authors: Niksoney A. Mendonça & Thaís E. Almeida — SciELO Preprints (preprint), 2025. Access preprint