Thaís Elias Almeida

A community-derived classification for extant lycophytes and ferns

Phylogeny has long informed pteridophyte classification. As our ability to infer evolutionary trees has improved, classifications aimed at recognizing natural groups have become increasingly predictive and stable. Here, we provide a modern, comprehensive classification for lycophytes and ferns, down to the genus level, utilizing a community‐based approach. We use monophyly as the primary criterion for the recognition of taxa, but also aim to preserve existing taxa and circumscriptions that are both widely accepted and consistent with our understanding of pteridophyte phylogeny. In total, this classification treats an estimated 11 916 species in 337 genera, 51 families, 14 orders, and two classes. This classification is not intended as the final word on lycophyte and fern taxonomy, but rather a summary statement of current hypotheses, derived from the best available data and shaped by those most familiar with the plants in question. We hope that it will serve as a resource for those wanting references to the recent literature on pteridophyte phylogeny and classification, a framework for guiding future investigations, and a stimulus to further discourse.

Diversity of ferns and lycophytes in Brazil

This compilation of ferns and lycophytes in Brazil is an update of the one published in 2010 in Catalogo de Plantas e Fungos do Brasil. The methodology consisted in collecting data from regional checklists, taxonomic revisions, and selected databases. Invited specialists improved the list accessing a website housed at the Jardim Botânico do Rio de Janeiro. The results show 1,253 species: 1,111 of ferns and 142 of lycophytes. This number is 6.5% higher than the previous one (1,176 spp.). The percentage of endemic species decreased from 38.2% to 36.7%. We recognized 36 families and 133 genera (vs. 33 families, 121 genera in 2010). The 10 most diverse families are Pteridaceae (196 spp.), Dryopteridaceae (179), Polypodiaceae (164), Hymenophyllaceae (90), Thelypteridaceae (86), Aspleniaceae (78), Lycopodiaceae (64), Selaginellaceae (55), Anemiaceae (51), and Cyatheaceae (45). The three most diverse genera are still Elaphoglossum (87 spp.), Thelypteris (85), and Asplenium (74). The richest phytogeographic domain continues to be in the Atlantic Rainforest with 883 species which also has the largest number of endemic and threatened species, followed by the Amazon Rainforest (503), Cerrado (269), Pantanal (30), Caatinga (26), and Pampa (eight). Minas Gerais remains as the richest state (657 spp. vs. 580 in 2010).

Pteridófitas do Parque Estadual do Jacupiranga, SP, Brasil

ABSTRACT – (Pteridophytes of Jacupiranga S tate Park, Sao Paulo S tate, Brazil). A floristic survey was carried out of the pteridophytesfrom Jacupiranga State Park, located in the municipal districts of Barra do Turvo, Cajati, Cananeia, Eldorado, Iporanga and Jacupiranga,in southern Sao Paulo state. Collecting was done in March and April/2005; voucher specimens are deposited in the BHCB, ESA, SPSFand UEC herbaria. Two hundred and twelve taxa (207 species, one subspecies and four varieties) distributed in 24 families and 65 generawere recorded. The most representative families were Thelypteridaceae with 23 species and three varieties and Polypodiaceae with 23species. The richest genera were Thelypteris with 22 species and three varieties and Asplenium with 18 species and one variety. Asregards life form, of the 212 taxa, 122 are terrestrial, 65 epiphytes, 17 lithophytes, three creepers and five with more than one habitat.The species Diplazium rieddelianum Kunze, Stigmatopteris ulei

A New Species of Microgramma (Polypodiaceae) from Brazil and Recircumscription of the Genus Based on Phylogenetic Evidence

Abstract We describe a new species of Microgramma (M. microsoroides) from the Atlantic Forest of Brazil, present a phylogeny of Microgramma, and provide a new generic circumscription. Microgramma comprises ca. 30 species in the Neotropics and perhaps two species in Africa and on islands in the Indian Ocean. Several species are contentious with regard to generic placement, and have been treated within Pleopeltis, Polypodium, and Solanopteris. From all known species of Microgramma, the new species differs by the sori forming irregular rows between adjacent primary veins. This finding is quite surprising, inasmuch as all other species in the genus have sori in two very strict rows, one row on each side of the midrib. The monophyly of Microgramma, as newly redefined, is strongly supported in chloroplast DNA-based phylogenetic analyses, using the genes rbcL and rps4, the rps4–trnS intergenic spacer, and the trnL–trnF intergenic spacer on 13 samples from 12 species. The new species is nested within Microgramma as sister to Microgramma megalophylla. Recognition of the relationships of M. microsoroides calls into question certain soral characters traditionally used to define genera of Polypodiaceae, as well as genera of ferns in other families. Sorus placement is notoriously variable in some genera and in many different families of Polypodiales.

Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera

Thelypteridaceae is one of the largest fern families, having about 950 species and a cosmopolitan distribution but with most species occurring in tropical and subtropical regions. Its generic classification remains controversial, with different authors recognizing from one up to 32 genera. Phylogenetic relationships within the family have not been exhaustively studied, but previous studies have confirmed the monophyly of the lineage. Thus far, sampling has been inadequate for establishing a robust hypothesis of infrafamilial relationships within the family. In order to understand phylogenetic relationships within Thelypteridaceae and thus to improve generic reclassification, we expand the molecular sampling, including new samples of Old World taxa and, especially, many additional neotropical representatives. We also explore the monophyly of exclusively or mostly neotropical genera Amauropelta, Goniopteris, Meniscium, and Steiropteris. Our sampling includes 68 taxa and 134 newly generated sequences from two plastid genomic regions (rps4-trnS and trnL-trnF), plus 73 rps4 and 72 trnL-trnF sequences from GenBank. These data resulted in a concatenated matrix of 1980 molecular characters for 149 taxa. The combined data set was analyzed using maximum parsimony and bayesian inference of phylogeny. Our results are consistent with the general topological structure found in previous studies, including two main lineages within the family: phegopteroid and thelypteroid. The thelypteroid lineage comprises two clades; one of these included the segregates Metathelypteris, Coryphopteris, and Amauropelta (including part of Parathelypteris), whereas the other comprises all segregates of Cyclosorus s.l., such as Goniopteris, Meniscium, and Steiropteris (including Thelypteris polypodioides, previously incertae sedis). The three mainly neotropical segregates were found to be monophyletic but nested in a broadly defined Cyclosorus. The fourth mainly neotropical segregate, Amauropelta, was found to include species considered to be part of Parathelypteris. In Old World thelypteroids, which correspond to nearly half the diversity in the family, an increase in sampling is still needed to resolve relationships and circumscription of genera, particularly in the christelloid clade (i.e., Amphineuron, Chingia, Christella, Pneumatopteris, Pronephrium, and Sphaerostephanos). Based on currently available knowledge, we propose the recognition of 16 genera in the family.

A comprehensive checklist of vascular epiphytes of the Atlantic Forest reveals outstanding endemic rates.

Abstract Knowledge of the geographic distribution of plants is essential to underpin the understanding of global biodiversity patterns. Vascular epiphytes are important components of diversity and functionality of Neotropical forests but, unlike their terrestrial counterparts, they are under-represented in large-scale diversity and biogeographic analyses. This is the case for the Atlantic Forest - one of the most diverse and threatened biomes worldwide. We provide the first comprehensive species list of Atlantic Forest vascular epiphytes; their endemism patterns and threatened species occurrence have also been analyzed. A list with 2,256 species of (hemi-)epiphytes - distributed in 240 genera and 33 families - is presented based on the updated Brazilian Flora Checklist. This represents more than 15% of the total vascular plant richness in the Atlantic Forest. Moreover, 256 species are included on the Brazilian Red List. More than 93% of the overall richness is concentrated in ten families, with 73% represented by Orchidaceae and Bromeliaceae species alone. A total of 78% of epiphytic species are endemic to the Atlantic Forest, in contrast to overall vascular plant endemism in this biome estimated at 57%. Among the non-endemics, 13% of epiphytic species also occur either in the Amazon or in the Cerrado - the other two largest biomes of Brazil – and only 8% are found in two or more Brazilian biomes. This pattern of endemism, in addition to available dated phylogenies of some genera, indicate the dominance of recent radiations of epiphytic groups in the Atlantic Forest, showing that the majority of divergences dating from the Pliocene onwards are similar to those that were recently reported for other Neotropical plants.

Two New Species of the Fern Genus Blechnum with Partially Anastomosing Veins from Northern Brazil

Abstract Two new species of Blechnum from Para State, northern Brazil, are described along with a diagnostic key and illustrations: Blechnum areolatum and B. longipilosum (Blechnaceae). Blechnum areolatum can be distinguished by: partially anastomosing veins, one or two pairs of pinnae and one- to two-celled hairs present only on the abaxial side of leaf blade. Blechnum longipilosum is characterized by: partially anastomosing veins, one to three pairs of reduced proximal pinnae and long hairs on rachises, costae, veins, laminar tissue, and indusia between sporangia. Both plants belong to the B. occidentale L. complex. Blechnum areolatum is similar to some specimens of B. lanceola Sw. with one pair of pinnae, but the veins in the latter species are completely free. Blechnum longipilosum has no known close relatives. The only Neotropical species so far known with partially anastomosing veins is B. heringeri Brade, which has erect rhizomes (vs. short creeping in B. longipilosum) and short, two- to three-celled hairs on leaf blades, which are deltoid and truncate at base. A key to the Neotropical species of Blechnum with partially anastomosing veins is provided.

New combinations in Neotropical Thelypteridaceae.

Abstract 288 new combinations of Neotropical Thelypteridaceae taxa are proposed in order to recognize monophyletic genera, based on the results of the most recent molecular phylogeny of the family, as well as the morphological uniformity of characters for each genus. The new nomenclatural combinations correspond to 186 Amauropelta taxa, 77 species of Goniopteris, and 25 Steiropteris taxa. A key to all native Neotropical genera of the family is also presented.

Global phylogeny and biogeography of the fern genus Ctenitis (Dryopteridaceae), with a focus on the Indian Ocean region

The diverse and pantropical genus Ctenitis, in the Dryopteridaceae, has been largely ignored in phylogenetic studies until now. In this study, we fill in this gap by reconstructing the first comprehensive phylogeny of the genus including 53 species currently recognized in the genus Ctenitis, among which seven species formerly were assigned to the genus Pseudotectaria and one to Heterogonium. Special emphasis was given to the sampling of species occurring in the African-Indian Ocean region. The presented results include reconstruction of a biogeographic scenario based on estimated divergence times and ancestral area reconstruction. Our findings confirm the inclusion, within Ctenitis, of the Indian Ocean species formerly placed in Pseudotectaria and Heterogonium. The crown group divergence was estimated to date back to the Oligocene or Early Miocene. The biogeographical scenario indicates an initial divergence of the Asian-Pacific ranges and the neotropical ranges, and a subsequent colonization of the Afro-Madagascan region by a lineage with neotropical ancestors. The Afro-Madagascan lineage splits into a lineage endemic to the Mascarene islands and a lineage occurring in Madagascar, the Comoros and Africa. The range expansion towards Africa and Madagascar was estimated to date back to the late Miocene, whereas the estimated ages for the onset of the diversification of the Mascarene diversity is consistent with the ages of these young, volcanic islands. The absence of any extant species of Ctenitis with a multi-continental distribution range and the rarity of inter-island dispersal and speciation in the Indian Ocean region suggest a limited contribution of long distance dispersal to the biogeographical history of this fern genus, versus a high contribution of local speciation.

Molecular phylogeny of the fern family Blechnaceae (Polypodiales) with a revised genus‐level treatment

Blechnaceae, a leptosporangiate fern family nested within eupolypods II, comprises 200–250 species, typically divided among seven to nine genera. Despite recent molecular studies of the family, it still lacks a modern taxonomic update based on broad sampling from the two centres of diversity—the Neotropics and Australasia/Oceania. To test generic circumscriptions, we have assembled the broadest dataset thus far, from three plastid regions (rbcL, rps4‐trnS, trnL‐trnF) and with taxonomic sampling focused on both major diversity centres. Our sampling includes 156 taxa and 178 newly generated sequences. We recognize three subfamilies, each corresponding to a highly supported clade across all analyses (maximum parsimony, Bayesian inference and maximum likelihood). The genera Salpichlaena, Stenochlaena and Telmatoblechnum are monophyletic, while Blechnum is polyphyletic, because Brainea, Doodia and Sadleria all nest within it. We outline and explain a plan to resolve the polyphyly of Blechnum by recognizing additional, monophyletic, segregate genera.