ABSTRACT

In the Municipality of Viçosa, one family, three genera, and five species of Gleicheniales occur: Dicranopteris flexuosa, Gleichenella pectinata, Sticherus bifidus, S. lanuginosus, and S. nigropaleaceus (Gleicheniaceae). All these species are very common in the area, forming dense thickets (“gleiquenais”) in disturbed areas, and co-occurring in these thickets. Dicranopteris flexuosa, G. pectinata, and S. bifidus are widespread in the Neotropics; Sticherus lanuginosus is widespread in South America and Costa Rica; and S. nigropaleaceous is endemic to the Brazilian Atlantic Forest. We provide nomenclatural corrections for S. lanuginosus. In Osmundales, only one species is found in Viçosa, occurring in marshes: Osmunda palustris (Osmundaceae). The taxonomy of Osmunda is still controversial, with the main Brazilian taxon named as O. palustris, O. regalis, O. regalis subsp. palustris, O. regalis var. spectabilis, O spectabilis, or O. spectabilis var. palustris. In our view, the best name for the taxon occurring in Viçosa is O. palustris, a taxon likely with a Neotropical distribution. Our taxonomic judgment is based on the re-interpretation of recent phylogenetic works, together with populational and morphological studies comparing Brazilian specimens with specimens from the U.S.A. We present descriptions, illustrations, examined specimens, taxonomic comments, and keys to identification.

Keywords:
gleiquenal; gleiquênia; Osmunda palustris; Osmunda regalis; Osmunda spectabilis; Sticherus

RESUMO

No município de Viçosa, uma família, três gêneros e cinco espécies de Gleicheniales ocorrem: Dicranopteris flexuosa, Gleichenella pectinata, Sticherus bifidus, S. lanuginosus e S. nigropaleaceus (Gleicheniaceae). Todas estas espécies são extremamente comuns na região, formando densos gleiquenais em áreas alteradas, e co-ocorrendo nestes gleiquenais. Dicranopteris flexuosa, G. pectinata e S. bifidus são amplamente distribuídos na região neotropical; Sticherus lanuginosus é amplamente distribuído na América do Sul e Costa Rica; e S. nigropaleaceous é endêmico à Floresta Atlântica brasileira. São providas correções nomenclaturais em S. lanuginosus. Em Osmundales, apenas uma espécie é encontrada em Viçosa, em ambientes paludosos: Osmunda palustris (Osmundaceae). A taxonomia de Osmunda é ainda controversa, na qual o principal táxon brasileiro vem sendo identificado como O. palustris, O. regalis, O. regalis subsp. palustris, O. regalis var. spectabilis, O. spectabilis, ou O. spectabilis var. palustris. Aqui, sugere-se que o melhor nome a ser adotado ao táxon ocorrente em Viçosa seja O. palustris, e ele provavelmente tenha distribuição neotropical. A decisão taxonômica foi baseada na re-interpretação de estudos filogenéticos recentes e em estudos populacionais e morfológicos, comparando espécimes brasileiros e norte-americanos. São apresentadas descrições, ilustrações, lista de material examinado, comentários taxonômicos e chaves de identificação.

Palavras-chave:
gleiquenal; gleiquênia; Osmunda palustris; Osmunda regalis; Osmunda spectabilis; Sticherus

Introduction

Gleicheniales and Osmundales are two ancient orders of ferns that arouse in the second radiation of the filicaleans, much before the modern Polypodiales. Both orders have fossils records starting on the Perminan, between 250 and 300 million years ago (Rothwell & Stockey 2008Rothwell, G.W. & Stockey, R.A. 2008. Phylogeny and evolution of ferns: a paleontological perspective. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. pp. 332-366.; Smith et al. 2008Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.). Gleicheniales is characterized by its species having generally long-creeping, underground rhizomes, which are protostelic or solenostelic, pseudo-dichotomous leaves, and sporangia with oblique annulus that are not very distinguishable from the capsule cells (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Smith et al. 2008Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.). Osmundales is characterized by massive stems with unique ectophloic solenosteles with rings of discrete xylem strands, hemi-dimorphic or dimorphic leaves, and large sporangia with lateral annulus (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Smith et al. 2008Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.).

Gleicheniales embraces three families (sensuPPG I, 2016PPG I (The Pteridophyte Phylogeny Group). 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603.): Dipteridaceae, with two genera, and 11 species; Gleicheniaceae, with six genera, and ca. 160 species; and Matoniaceae, with two genera, and four species. Among these families, only Gleicheniaceae occurs in Brazil, represented by Dicranopteris (with three species), Gleichenella (one sp.), and Sticherus (10-15 spp.) (Gonzales & Kessler 2011Gonzales, J. & Kessler, M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1-54.; Lima & Salino 2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234., 2020Lima, L.V. & Salino, A. 2020. Gleicheniaceae. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91158 . (Access in 29-VIII-2022).
http://floradobrasil.jbrj.gov.br/reflora... ). The genus name Gleichenia has been and is sometimes applied to Brazilian plants (e.g., Lisbôa 1956Lisbôa, M.A. 1956. Ptéridophytes de Ouro Preto. Anais da Escola de Minas de Ouro Preto 29: 21-77.; Sehnem 1970Sehnem A. 1970. Gleiqueniáceas. In: R. Reitz (ed.) Flora Ilustrada Catarinense. Itajaí, Instituto Anchietano de Pesquisas, São Leopoldo.), but this genus is Paleotropical. On the other hand, common names in Portuguese, such as “gleiquênia” and “gleiquenal” (= Gleicheniaceae thickets), are correctly used for these plants. In Brazil, such plants are very common, forming dense thickets in disturbed, open areas (e.g., Sehnem 1970Sehnem A. 1970. Gleiqueniáceas. In: R. Reitz (ed.) Flora Ilustrada Catarinense. Itajaí, Instituto Anchietano de Pesquisas, São Leopoldo.; Windisch 1994Windisch, P.G. 1994. Pteridófitas do Estado de Mato Grosso: Gleicheniaceae. Bradea 6: 304-311.; Prado 2004aPrado, J. 2004a. Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP. Pteridophyta: 8. Gleicheniaceae. Hoehnea 31: 33-37.; Schwartsburd & Labiak 2007Schwartsburd, P.B. & Labiak, P.H. 2007. Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34(2): 159-209.; Lima & Salino 2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.). The ecological niche of pioneer plants occurring in disturbed, sunny areas with poor soils, is common across the family (Kessler & Smith 2018Kessler, M. & Smith, A.R. 2018. Prodomus of a fern Flora for Bolivia. XI. Gleicheniaceae. Phytotaxa, 344: 53-63.), and it is resulted from their habit allied to phytotoxic compounds they produce (Østergaard-Andersen & Ølgaard, 2001Østergaard-Andersen, E & Øllgaard, B. 2001. Gleicheniaceae. In: G. Harling & L. Andersson (eds.). Flora of Ecuador 66. Göteborg University, Göteborg. pp. 107-169.). Gleichenella pectinata, for example, produces compounds that are allelopatic to many other plants (Muller et al. 2007Muller, C., Chagas, F.F., Peres, T.L.P., Hess, S.C., Faccenda, O & Daloso, D.M. 2007. Potencial fitotóxico de algumas espécies de Gleicheniaceae sobre Allium cepa L. Revista Brasileira de Biociências 5: 45-47.), and with insecticide action (Moliterno & Abreu 2016Moliterno, A.A.C. & Abreu, P.F. 2016. Atividade inseticida dos extratos aquosos de Gleichenella pectinata (Willd) Ching (Gleicheniaceae) contra Atta laevigata (Smith, 1858) (Hymenoptera: Formicidae). CES Revista 30: 56-68.).

Gleicheniaceae, as currently circumscribed (e.g.,PPG I 2016PPG I (The Pteridophyte Phylogeny Group). 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603.) is monophyletic, but some genera sill require new and modern circumscriptions: the Old Word genus Gleichenia has shown to be polyphyletic (Perrie et al. 2007Perrie, L.R., Bayly, M.J., Lehnbach, C.E. & Brownsey, P.J. 2007. Molecular phylogenetics and molecular dating of the New Zealand Gleicheniaceae. Brittonia 59: 129-141.; Li et al. 2010Li C, Lu S, Ma J, Yang Q. 2010. Phylogeny and divergence of Gleicheniaceae inferred from three plastid genes. Acta Palaeontologica Sinica 49: 64-72.; Wei et al. 2021Wei, Z., Xia, Z., Shu, J., Shang, H., Maxwell, S.J., Chen, L., Zhou, X., Xi, W., Adjie, B., Yuan, Q., Cao, J. & Yan, Y. 2021. Phylogeny and taxonomy on cryptic species of forked ferns of Asia. Frontiers in Plant Science 12: 748562.), and the Pantropical Sticherus has shown to be paraphyletic (Lima et al. 2023Lima, L.V., Salino, A., Kessler, M., Rouhan, G., Testo, W.L., Argolo, C.S., GoFlag Consortium & Almeida, T.E. 2023. Phylogenomic evolutionary insights in the fern family Gleicheniaceae. Molecular Phylogenetics and Evolutions 184: 107782.). According to Lima et al. (2023Lima, L.V., Salino, A., Kessler, M., Rouhan, G., Testo, W.L., Argolo, C.S., GoFlag Consortium & Almeida, T.E. 2023. Phylogenomic evolutionary insights in the fern family Gleicheniaceae. Molecular Phylogenetics and Evolutions 184: 107782.), Dicranopteris and Gleichenella are monophyletic, and the Brazilian species of Sticherus form a phylogenetic group along with the Neotropical remaining species of Sticherus. Further studies should review whether the Brazilian species should be kept in Sticherus s. str., or if they should be combined into another genus name.

The number of Brazilian species of Sticherus is still controversial, due to two conflicting revisionary works on the genus: while Gonzales & Kessler (2011Gonzales, J. & Kessler, M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1-54.) recognized 12 species in Brazil (four of which occurring in Minas Gerais), Lima & Salino (2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.) recognized 11 species (seven in Minas Gerais). At first sight, 11 and 12 species seem similar, but a detailed look at their works shows there are many differences on which species names are accepted and which are synonymized. Later, Lima & Salino (2020Lima, L.V. & Salino, A. 2020. Gleicheniaceae. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91158 . (Access in 29-VIII-2022).
http://floradobrasil.jbrj.gov.br/reflora... ) also recognized 12 species.

Osmundales comprises solely family Osmundaceae, with four extant genera and about 20 extant species (PPG I 2016PPG I (The Pteridophyte Phylogeny Group). 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603.): Claytosmunda (one species), Leptopteris (six spp.), Osmunda (ca. four spp.), Osmundastrum (one sp.), Plenasium (ca. four spp.), and Todea (two spp.). Phylogenetically, Osmundales plays an important role as a link between the eusporangiate and the leptosporangiate ferns: they have leptoesporangia, but formed by more than one initial cell, and enormous in size, when compared to other leptoesporangia (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Smith et al. 2008Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.).

In Brazil, Osmundaceae is represented by Osmunda and Osmundastrum (Sylvestre et al. 2020Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2020. Osmundaceae. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91530 (Accessed in: 29-VIII-2022).
http://floradobrasil.jbrj.gov.br/reflora... , 2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.). Osmundastrum is represented solely by the sub-comopolitan Osmundastrum cinnamomeum (L.) C. Presl. The species of Osmunda, on the other hand, are still debatable. Many authors considered two species in Brazil: one widespread and another endemic to the Cerrado region and southern Brazil. The endemic species has been commonly named as Osmunda gracilis Link (by Sturm 1859Sturm, J.G. 1859. Ophioglosseae, Marattiaceae, Osmundaceae, Schizaeaceae, Gleicheniaceae ey Hymenophylleae. In: C.F.P. Martius & A.G. Eichler. Flora Brasiliensis. Monachii: Lipsiae apud Frid. Fleischer in comm., Vol. 1, Part 2.; Fée 1869Fée, A.L.A. 1869. Cryptogames Vasculaires (fougères, lycopodiacées, hydroptéridées, équisétacées) du Brésil. J.B. Baillière et fils, Libraires, Paris.; Sehnem 1967Sehnem A. 1967. Osmundáceas. In: R. Reitz (ed.) Flora Ilustrada Catarinense. Itajaí, Instituto Anchietano de Pesquisas, São Leopoldo.; Arana & Ponce 2015Arana, M.D. & Ponce, M.M. 2015. Osmundaceae en Argentina, Paraguay y Uruguay. Darwiniana 3: 27-37.; Sylvestre et al. 2020Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2020. Osmundaceae. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91530 (Accessed in: 29-VIII-2022).
http://floradobrasil.jbrj.gov.br/reflora... ), but recently as Osmunda piresii Brade (Sylvestre et al. 2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.). The widespread taxon has been treated either as a cosmopolitan species, Osmunda regalis L. (e.g.,Lisbôa 1956Lisbôa, M.A. 1956. Ptéridophytes de Ouro Preto. Anais da Escola de Minas de Ouro Preto 29: 21-77.; Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Schwartsburd & Labiak 2007Schwartsburd, P.B. & Labiak, P.H. 2007. Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34(2): 159-209.), as an American variety, Osmunda regalis var. spectabilis (Willd.) A. Gray (e.g.,Prado 2004bPrado J. 2004b. Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP. Pteridophyta: 15. Osmundaceae. Hoehnea 31: 93-96.), as an American species, Osmunda spectabilis Willd. (e.g.,Fée 1869Fée, A.L.A. 1869. Cryptogames Vasculaires (fougères, lycopodiacées, hydroptéridées, équisétacées) du Brésil. J.B. Baillière et fils, Libraires, Paris.; Arana & Ponce 2015Arana, M.D. & Ponce, M.M. 2015. Osmundaceae en Argentina, Paraguay y Uruguay. Darwiniana 3: 27-37.; Sylvestre et al. 2020Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2020. Osmundaceae. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91530 (Accessed in: 29-VIII-2022).
http://floradobrasil.jbrj.gov.br/reflora... , 2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.), as a Neotropical variety, Osmunda spectabilis var. palustris (Schrad.) C. Presl (e.g.,Presl 1845Presl, C.B. 1845. Supplementum tentaminis pteridographiae. Aulica Filiorum Amadei Haase, Prague.), as a Brazilian subspecies, Osmunda regalis subsp. Palustris (Schrad.) Á. Löve & D. Löve (e.g., Löve & Löve 1977Löve, Á. & Löve, D. 1977. New combinations in ferns. Taxon 26: 324-326.), or as a Brazilian species, Osmunda palustris Schrad. (e.g.,Sturm 1859Sturm, J.G. 1859. Ophioglosseae, Marattiaceae, Osmundaceae, Schizaeaceae, Gleicheniaceae ey Hymenophylleae. In: C.F.P. Martius & A.G. Eichler. Flora Brasiliensis. Monachii: Lipsiae apud Frid. Fleischer in comm., Vol. 1, Part 2.; Sehnem 1967Sehnem A. 1967. Osmundáceas. In: R. Reitz (ed.) Flora Ilustrada Catarinense. Itajaí, Instituto Anchietano de Pesquisas, São Leopoldo.).

Phylogenetic studies show that Osmunda regalis s.l., considered as a cosmopolitan species, is artificial, rendering a paraphyletic species (Metzgar et al. 2008Metzgar, J.S., Skog, J.E., Zimmer, E.A. & Pryer, K.M. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany 33: 31-36.; Tsutsumi et al. 2021Tsutsumi, C., Yatabe-Kakugawa, Y., Hirayama, Y., Chiou, W.-L. & Kato, M. 2021. Molecular analyses of the disjunctly distributed Osmunda regalis and O. japonica (Osmundaceae), with particular reference to introgression and hybridization. Plant Systematics and Evolution 307: 35.). Rather, there are several monophyletic clades within Osmunda, including cryptic species of the Osmunda regalis-complex, well-defined species, as well as hybrids. Seven clades were presented by Tsutsumi et al. (2021Tsutsumi, C., Yatabe-Kakugawa, Y., Hirayama, Y., Chiou, W.-L. & Kato, M. 2021. Molecular analyses of the disjunctly distributed Osmunda regalis and O. japonica (Osmundaceae), with particular reference to introgression and hybridization. Plant Systematics and Evolution 307: 35.): 1. Containing O. japonica Thunb. And O. lancea Thunb.; 2. Containing only O. lancea; 3. Containing “O. regalis” from the Americas; 4. Containing O. regalis from Europe; 5. Containing “O. regalis” from Cameroon; 6. Containing “O. regalis” from Madagascar; and 7. Containing “O. regalis” from India and O. hybrida Tsutsumi, S. Matsumoto, Y. Yatabe, Y. Hiray. & M. Kato. The type of O. regalis is European, therefore the European clade retains the name O. regalis s. str. Judging the American clade as a monophyletic species, Sylvestre et al. (2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.) adopted the name O. spectabilis for the species widespread in the Americas.

Continuing the papers on the Fern Flora of Viçosa, in which Medeiros & Schwartsburd (in pressMedeiros, M.T. & Schwartsburd, P.B. in press. Fern Flora of Viçosa, Minas Gerais State, Brazil: Blechnaceae (Polypodiopsida). Hoehnea: in press.) presented the Blechnaceae, Pena & Schwartsburd (2022Pena, N.T.L. & Schwartsburd, P.B. 2022. Fern flora of Viçosa, Minas Gerais State, Brazil: Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae. Hoehnea 50: e152022.), the Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae, Matos & Schwartsburd (2022Matos, F.B. & Schwartsburd, P.B. 2022. Ferns of Viçosa, Minas Gerais, Brazil: Didymochlaenaceae and Dryopteridaceae. Rodriguésia 73: e00282022.), the Dryopteridaceae, Gonçalves da Silva & Schwartsburd (2017Gonçalves da Silva, A. & Schwartsburd, P.B. 2017. Ferns of Viçosa, Minas Gerais State, Brazil: Polypodiaceae (Polypodiales, Filicopsida, Tracheophyta). Hoehnea 44: 251-268.), the Polypodiaceae, Miranda & Schwartsburd (2017Miranda, C.V. & Schwartsburd, P.B. 2016. Aquatic ferns from Viçosa (MG, Brazil): Salviniales (Filicopsida; Tracheophyta). Brazilian Journal of Botany 39: 935-942.), the Salviniales (Marsileaceae and Salviniaceae), and Rabelo & Schwartsburd (2016Rabelo, L.S. & Schwartsburd, P.B. 2016. Ferns of Viçosa, MG, Brazil: Schizaeales (Filicopsida, Tracheophyta), with special reference to hybrids. Brittonia 68: 379-396.), the Schizaeales (Anemiaceae and Lygodiaceae), we here present the taxonomic treatment for Gleicheniales (Gleicheniaceae) and Osmundales (Osmundaceae). So far, these publications have unveiled the occurrence of 64 species and seven hybrids for the local Flora. Among these taxa, 25 are endemic to the Brazilian Atlantic Forest, and among them, one species, one variety and three hybrids have been described as new to science. In addition, six names have been rescued from recent synonymy (including the proposal of one new combination), and three hybrids have been newly recorded for the Brazilian territory.

Material and methods

We have been collecting ferns in Viçosa since July of 2012, especially in the protected areas, such as Estação Mata do Paraíso, Recanto das Cigarras, Mata do Seu Nico, and smaller forest fragments. In the case of Gleicheniales, on the other hand, the disturbed areas are the best place to find them. And so, our main collections are from disturbed areas, alongside roads, and forest clearings. The forest fragments of Viçosa are considered Semi-Decidual Seasonal Forest (IBGE 2012IBGE. 2012. Manual técnico da vegetação brasileira. Instituto Brasileiro de Geografia e Estatística, 2a ed., Rio de Janeiro. ), and they range from around 600 to 900 m altitude (Gonçalves da Silva & Schwartsburd 2017Gonçalves da Silva, A. & Schwartsburd, P.B. 2017. Ferns of Viçosa, Minas Gerais State, Brazil: Polypodiaceae (Polypodiales, Filicopsida, Tracheophyta). Hoehnea 44: 251-268.). We collected specimens following usual procedures for ferns (e.g.,Fidalgo & Bononi 1989Fidalgo, O. & Bononi, V.L.R. 1989. Técnicas de coleta, preservação e herborização de material botânico. Instituto de Botânica, São Paulo.).

We observed the ferns under magnifying glass to see hairs, scales, grooves, and veins, and prepared specimens and incorporated them into herbarium VIC, with duplicates sent to UCS (and others to be sent to RB and SP). We contributed with 73 herbarium specimens. We analyzed the fern collection of VIC, in which another 25 specimens were studied. We also consulted herbaria online, to search for additional specimens from Viçosa, especially those collected by Y. Mexia and J.G. Kuhlmann, from the following websites: SpeciesLink (http://www.splink.org.br/), Reflora (http://floradobrasil.jbrj.gov.br/reflora/herbarioVirtual/), Jstor Global Plants (https://plants.jstor.org/), and the Pteridophyte Collections Consortium (http://www.pteridoportal.org/portal/collections/). Another 24 specimens were studied online. Morphological terms follow Lellinger (2002Lellinger, D.B. 2002. A modern multilingual glossary for taxonomic pteridology. Pteridologia 3: 1-263.). The illustrations were prepared by Reinaldo Pinto.

Specifically, for Osmundaceae, one of us (Schwartsburd) have collected and studied populations in the field in many places in Brazil (in the States of Minas Gerais, Espírito Santo, and Paraná), in the USA (State of Vermont), and in Italy.

Results and discussion

Diversity and Distribution - From Viçosa and surroundings, we report the occurrence of one family, three genera, and five species of Gleicheniales: Dicranopteris flexuosa, Gleichenella pectinata, Sticherus bifidus, S. lanuginosus, and S. nigropaleaceus (Gleicheniaceae). All these species are extremely common in the area, forming dense thickets (“gleiquenais”) in disturbed areas, and co-occurring in these thickets. The slopes (“barrancos”) around Viçosa are either tamed by the Gleicheniaceae, either by exotic grasses (“braquiária”, “capim-gordura”, etc.). There is an apparent competition between the group that will form the thicket.

Dicranopteris flexuosa, Gleichenella pectinata, and Sticherus bifidus are widespread in the Neotropics; Sticherus lanuginosus is widespread in South America; and S. nigropaleaceous is endemic to the Brazilian Atlantic Forest. The three Sticherus spp. from Viçosa are considered valid by both the treatments of Gonzales & Kessler (2011Gonzales, J. & Kessler, M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1-54.) and Lima & Salino (2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234., 2020).

For Osmundales, only one species is found in Viçosa, occurring exclusively on marshes, on flooded soils: Osmunda palustris (Osmundaceae). The taxonomy of Osmunda is still controversial, with the main Brazilian taxon identified as either O. palustris (e.g.,Sturm 1859Sturm, J.G. 1859. Ophioglosseae, Marattiaceae, Osmundaceae, Schizaeaceae, Gleicheniaceae ey Hymenophylleae. In: C.F.P. Martius & A.G. Eichler. Flora Brasiliensis. Monachii: Lipsiae apud Frid. Fleischer in comm., Vol. 1, Part 2.; Sehnem 1967Sehnem A. 1967. Osmundáceas. In: R. Reitz (ed.) Flora Ilustrada Catarinense. Itajaí, Instituto Anchietano de Pesquisas, São Leopoldo.), O. regalis (e.g.,Lisbôa 1956Lisbôa, M.A. 1956. Ptéridophytes de Ouro Preto. Anais da Escola de Minas de Ouro Preto 29: 21-77.; Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Schwartsburd & Labiak 2007Schwartsburd, P.B. & Labiak, P.H. 2007. Pteridófitas do Parque Estadual de Vila Velha, Ponta Grossa, Paraná, Brasil. Hoehnea 34(2): 159-209.), O. regalis var. spectabilis (e.g., Prado 2004Prado J. 2004b. Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP. Pteridophyta: 15. Osmundaceae. Hoehnea 31: 93-96.b), O. regalis subsp. palustris (e.g.,Löve & Löve 1977Löve, Á. & Löve, D. 1977. New combinations in ferns. Taxon 26: 324-326.), O. spectabilis (e.g.,Fée 1869Fée, A.L.A. 1869. Cryptogames Vasculaires (fougères, lycopodiacées, hydroptéridées, équisétacées) du Brésil. J.B. Baillière et fils, Libraires, Paris.; Arana & Ponce 2015Arana, M.D. & Ponce, M.M. 2015. Osmundaceae en Argentina, Paraguay y Uruguay. Darwiniana 3: 27-37.; Sylvestre et al. 2020Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2020. Osmundaceae. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91530 (Accessed in: 29-VIII-2022).
http://floradobrasil.jbrj.gov.br/reflora... , 2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.), or O. spectabilis var. palustris (e.g.,Presl 1845Presl, C.B. 1845. Supplementum tentaminis pteridographiae. Aulica Filiorum Amadei Haase, Prague.).Tsutsumi et al. (2021Tsutsumi, C., Yatabe-Kakugawa, Y., Hirayama, Y., Chiou, W.-L. & Kato, M. 2021. Molecular analyses of the disjunctly distributed Osmunda regalis and O. japonica (Osmundaceae), with particular reference to introgression and hybridization. Plant Systematics and Evolution 307: 35.) clearly showed that Osmuda regalis, considered as a sub-cosmopolitan species, is paraphyletic. Rather, to reach a monophyletic classification, the geographical clades of O. regalis s.l. should be regarded as different species. Osmunda regalis s. str. is kept for the European species, since its type is European. Tsutsumi et al. (2021Tsutsumi, C., Yatabe-Kakugawa, Y., Hirayama, Y., Chiou, W.-L. & Kato, M. 2021. Molecular analyses of the disjunctly distributed Osmunda regalis and O. japonica (Osmundaceae), with particular reference to introgression and hybridization. Plant Systematics and Evolution 307: 35.: Figure 5) presented a clade formed by populations from the Americas, which they named as the “American lineage” - this led Sylvestre et al. (2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.), for example, to name the widespread “species” from the Americas as O. spectabilis, whose type is from the USA. On the other hand, the lack of samples from the Neotropics masked the formation of two sub-clades clearly segregated within the American clade: one formed by the five samples of Osmunda from the USA, and another formed by a sample from Mexico and a sample from Misiones, Argentina, in the border with Brazil and Paraguay.

After our studies on living populations and morphological comparisons between Osmunda populations from Brazil (from Minas Gerais, Espírito Santo, and Paraná) and two populations from the USA (Vermont), we reached the conclusion that they represent two very distinct entities that we would regard as two different species: Osmunda palustris from Brazil (whose type is from Brazil), probably occurring in the whole Neotropics, and O. spectabilis s. str. occurring in the USA (and possibly in Canada). This conception is similar to those of Sturm (1859Sturm, J.G. 1859. Ophioglosseae, Marattiaceae, Osmundaceae, Schizaeaceae, Gleicheniaceae ey Hymenophylleae. In: C.F.P. Martius & A.G. Eichler. Flora Brasiliensis. Monachii: Lipsiae apud Frid. Fleischer in comm., Vol. 1, Part 2.) and Sehnem (1967Sehnem A. 1967. Osmundáceas. In: R. Reitz (ed.) Flora Ilustrada Catarinense. Itajaí, Instituto Anchietano de Pesquisas, São Leopoldo.) for the adoption of the species level to the Brazilian (Neotropical) taxon, and also especially similar to that of Presl (1845Presl, C.B. 1845. Supplementum tentaminis pteridographiae. Aulica Filiorum Amadei Haase, Prague.), who pointed out differences between the US/Canadian O. spectabilis var. spectabilis and the Neotropical O. spectabilis var. palustris (for which he cited specimens from Mexico and Brazil). Our judgment is based on a different interpretation of the phylogenetic hypothesis presented by Tsutsumi et al. (2021Tsutsumi, C., Yatabe-Kakugawa, Y., Hirayama, Y., Chiou, W.-L. & Kato, M. 2021. Molecular analyses of the disjunctly distributed Osmunda regalis and O. japonica (Osmundaceae), with particular reference to introgression and hybridization. Plant Systematics and Evolution 307: 35.), on the field studies and collections of populations of Osmunda spp. in several places in Brazil, in the USA, and in Italy, and in detailed morphological comparisons between them. This is a primary hypothesis, which, may be further investigate in the future with the addition of Brazilian samples (and other Neotropical as well) to the global phylogeny of Osmunda - which, in fact, we are starting to perform.

Regarding the habit and habitat differences, Osmunda palustris differs from O. spectabilis by living in tropical to sub-tropical habitats and being evergreen, whereas O. spectabilis lives on temperate habitats, commonly under the snow in some months, in which they loose all their leaves. The leaves of both species are different in size as well, with those of O. palustris reaching only 70-100 cm long (to 120 cm maximum, in exceptional cases - Figure 4a ), whereas O. spectabilis has longer leaves, commonly 100-150 cm long.

Morphologically, in addition to the differences in leaf sizes, Osmunda palustris further differs from O. spectabilis by the chartaceous to coriaceous laminae (vs. membranaceous), by the pinnae tappering at the apexes, producing apical pinnules ca. 1/3 the length of the lateral pinnules (vs. pinnae with conform apexes; the apical pinnules about the same length of the lateral ones), by the opposite to sub-opposite lateral pinnules (vs. alternate), and by the strongly raised veins (vs. immersed to slightly raised) (Figures 4a-cvs.Figures 4f-g ) . Some of these differences have already been pointed out by Tryon & Tryon (1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.), but they did not consider the two taxa as distinct, and by Presl (1845Presl, C.B. 1845. Supplementum tentaminis pteridographiae. Aulica Filiorum Amadei Haase, Prague.), who considered them as two different varieties.

Taxonomic Treatment

Gleicheniales

Rhizomes protostelic, solenostelic, or with apering solenosteles; leaves monomorphic or dimorphic; sporangia generally with oblique annuli, or rarely sub-vertical (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Smith et al. 2006Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.).

Gleicheniaceae C. Presl

Plants terrestrial or epipetric, commonly forming thickets. Rhizomes long-creeping, protostelic or rarely solenostelic; leaves monomorphic, indeterminate, generally pseudo-dichotomous, with buds at the furcations; veins free; sori abaxial, exsindusiate; sporangia with oblique annuli; spores trilete or monolete (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Smith et al. 2006Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.; Lima & Salino 2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.).

Six genera: Dicranopteris, Diplopterygium (not occurring in Brazil), Gleichenella, Gleichenia (not occurring in Brazil), Sticherus, and Stromatopteris (not occurring in Brazil).

Key to genera of Gleicheniaceae from Viçosa

1. Rhizomes and laminar buds with scales ……………..……………………………………….. Sticherus

1. Rhizomes and laminar buds with hairs

2. Laminae isotomically branched; aphlebioid pinnules present at the base of furcations ………………………………………...………...…...…………………………. Dicranopteris

2. Laminae anisotomically branched; aphlebioid pinnules absent ……………..…… Gleichenella

Dicranopteris Bernh.

Plants terrestrial or epipetric; rhizomes with hairs; leaves scandent; laminae isotomically branched; aphlebioid pinnules present at the base of furcations; buds at the base of furcations covered with hairs; segments abaxially glabrous or with hairs; spores trilete (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Lima & Salino 2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.).

Dicranopteris flexuosa (Schrad.) Underw., Bull. Torrey Bot. Club 34(5): 254. 1907. Mertensia flexuosa Schrad., Gött. Gel. Anz. 863. 1824. Gleichenia flexuosa (Schrad.) Mett., Ann. Mus. Bot. Lugduno-Batavi 1: 50. 1863.

Figures 1a-f

Plants terrestrial or rupestrial, thicket-forming. Rhizomes stramineous, 2.5-3.6 mm diam.; hairs reddish-brown. Leaves scrambling, to 3 m long; stipes to 1 m long, 2.5-3.1 mm diam., greenish to stramineous, glabrous; laminae 1-5-furcate, isotomically divided; rachises glabrous; accessory leaflets present at pinnae bases and subsequent divisions; buds with stiff, reddish-brown hairs; ultimate segments linear; midveins abaxially with sparse, reddish-brown hairs, adaxially glabrous; veins 1-2-furcate, abaxially with globose, orange, glandular hairs, adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; laminar margins flat.

Distribution: Widespread in the Neotropics. In Viçosa it is found on road-sides, disturbed areas, forest edges and ravines, under full sunlight exposure.

Material examined: BRAZIL. Minas Gerais: Viçosa, Belvedere, 11-IX-1977, “Rosane & Rosângela” s.n. (VIC-7931); Viçosa, Belvedere, 11-IX-1977, “Rosane & Rosângela” s.n. (VIC-7938); Viçosa, Belvedere, 11-IX-1977, “Rosane & Rosângela” s.n. (VIC-7939); Viçosa, Campus da UFV, final da mata da Biologia, 750 m, 16-VIII-2013, P.B. Schwartsburd et al. 2827 (VIC); Viçosa, Universidade Federal de Viçosa, ao final da PH Rolfs, próximo ao condomínio novo, 2-VI-2016, P.B. Schwarstsburd et al. 3781 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, ao final da PH Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwarstsburd & N.S. Smith-Braga 3797 (VIC); Viçosa, campus da UFV, 31-III-2019, V.S. Miranda et al. 74 (UCS, VIC); Viçosa, campus da UFV, 31-III-2019, V.S. Miranda et al. 75 (VIC); Viçosa, campus da UFV, 31-III-2019, V.S. Miranda et al. 76 (VIC); Viçosa, campus da UFV, 31-III-2019, V.S. Miranda et al. 77 (VIC); Viçosa, campus da UFV, em frente ao Herbário, 31-III-2019, V.S. Miranda et al. 79 (VIC); Viçosa, Mata do Paraíso, 24-IX-2019, V.S. Miranda et al. 90 (VIC).

Dicranopteris flexuosa is easily recognized by the isotomically divided laminae with accessory leaflets at the base of pinnae and next branches, and thus differing from Gleichenella pectinata, which has anisotomically divided laminae without accessory leaflets. Both D. flexuosa and G. pectinata are thicket forming, differing from Sticherus spp., which do not form thickets, but grow within D. flexuosa and G. pectinata thickets (gleiquenais).

Gleichenella Ching

Plants terrestrial; rhizomes with hairs; leaves scandent; laminae anisotomically branched; aphlebioid pinnules absent; buds at the base of furcations covered with hairs; segments abaxially glabrous or with hairs; spores monolete (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Lima & Salino 2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.).

Gleichenella pectinata (Willd.) Ching, Sunyatsenia 5(4): 276. 1940. Mertensia pectinata Willd., Kongl. Vetensk. Acad. Handl. 25: 168, t. 4. 1804. Gleichenia pectinata (Willd.) C. Presl, Reliq. Haenk. 1(1): 71. 1825. Dicranopteris pectinata (Willd.) Underw., Bull. Torrey Bot. Club 34(5): 260. 1907.

Figures 1g-j

Plants terrestrial, thicket-forming. Rhizomes stramineous to brown, 3.7-4.8 mm diam; hairs stiff, brown. Leaves scrambling, to 3 m long; petioles to 1.5 m long, 3.7-4 mm diam, greenish to stramineous; laminae 1-4-furcate, anisotomically divided; rachises glabrous; accessory leaflets absent; buds with stiff, brown hairs; ultimate segments linear; midveins abaxially with stellate hairs, adaxially glabrous; veins 1-2-furcate, abaxially with stellate hairs, adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; laminar margins revolute.

Distribution: Widespread in the Neotropics. In Viçosa it is found on road-sides, disturbed areas, forest edges and ravines, under full sunlight exposure.

Material examined: BRAZIL. Minas Gerais: Viçosa, ESAV, 24-XII-1958, H.S. Irwin 2308 (VIC); Viçosa, Campus da UFV, pastagem da Zootecnia, 20-V-1997 S.J.S. Neto s.n. (UCS, VIC-22923); Viçosa, Campus da UFV, Mata da Biologia, Barranco entre Belvedere I e Belvedere das Bandeiras, 10-IX-1998, G.E. Valente & L.A. Basílio 352 (VIC); Viçosa, Mata da Biologia, Recanto das Cigarras, primeira praça, 10-IX-1998, G.E. Valente & L.A. Basílio 354 (VIC); Viçosa, Mata da Prefeitura, 26-VII-1999, A.F. Carvalho 704 (VIC); Viçosa, Campus da UFV, final da Mata da Biologia, estrada 20º45’31”S, 42º51’56”W, 750 m, 16-VIII-2013, P.B. Schwartsburd et al. 2827b (VIC); Viçosa, Campus da UFV, Avenida P.H. Rolfs, Próximo ao R.U. 2, 2-VI-2014, P.B. Schwartsburd & T. Campos 3265 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, ao final da PH Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwarstsburd et al. 3778 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, Estrada para a criação de Equídeos, 13-X-2016, P.B. Schwartsburd & N.S. Smith-Braga 3798 (UCS, VIC); Viçosa, Mata da Biologia, Recanto das Cigarras entre as churrasqueiras e Bandeiras, esquina da rotatória, 20°45’18”S, 42°10’40”W, 28-III-2019, V.S. Miranda et al. 56 (UCS, VIC); Viçosa, Mata da Biologia, Recanto das Cigarras entre as churrasqueiras e Bandeiras, esquina da rotatória, 20°45’18”S, 42°10’40”W, 28-III-2019, V.S. Miranda et al. 59 (UCS, VIC); Viçosa, Mata da Biologia, Recanto das Cigarras, 29-III-2019, V.S. Miranda et al. 66 (UCS, VIC).

See the comments of Dicranopteris flexuosa for comparison with Gleichella pectinata.

Sticherus C. Presl

Plants terrestrial or epipetric; rhizomes with scales; leaves erect or scandent; laminae isotomically branched; aphlebioid pinnules absent; buds at the base of furcations covered with scales; segments abaxially glabrous or with scales; spores monolete (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Gonzales & Kessler 2011Gonzales, J. & Kessler, M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1-54.; Lima & Salino 2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.).

Key to species of Sticherus from Viçosa

1. Leaves erect; ultimate segments about two times longer than wider; bud scales without blackish stains ……………………………………..…………………………………………. S. lanuginosus

1. Leaves prostrate or scandent; ultimate segments about four times longer than wider; bud scales entirely black or with black stains

2. Leaves scandent; bud scales entirely black; laminar tissue between the veins abaxially glabrous ………………………………………………………..…………………..... S. nigropaleaceous

2. Leaves prostrate; bud scales stramineous with blackish stains; laminar tissue between the veins abaxially lanose, with arachnoid scales .……………………………..…………………. S. bifidus

Sticherus bifidus (Willd.) Ching, Sunyatsenia 5(4): 282. 1940. Mertensia bifida Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 168, t. 5, fig. B. 1804. Gleichenia bifida (Willd.) Spreng., Syst. Veg. ed. 16 [Sprengel] 4(1): 27. 1827. Dicranopteris bifida (Willd.) Maxon, N. Amer. Fl. 16(1): 60. 1909.

Figures 2a-f

Plants terrestrial. Rhizomes brown, 2.2-3.3 mm diam; scales brown, laceate, rigid, with dentate margins. Leaves prostrate; stipes 20-40 cm long, 2.0-2.7 mm diam., greenish to brownish, with stramineous scales with fimbriate margins; rachises terete, with stramineous scales with blackish stains, with fimbriate margins; buds with stramineous scales with blackish stains; ultimate segments linear, about four times longer than wider; midveins abaxially with fimbriate scales and arachnoid scales, adaxially glabrous; veins 1-furcate, abaxially with fimbriate scales and arachnoid scales, adaxially glabrous; laminar tissue between the veins abaxially lanose, adaxially glabrous; laminar margins flat to revolute.

Distribution: Widespread in the Neotropics. In Viçosa it is found on road-sides, disturbed areas, forest edges and ravines, under full or part-time sunlight exposure.

Material examined: BRAZIL. Minas Gerais: Porto Firme, 10 km da estrada de Viçosa, Sítio São Domingos, 25-I-2014, A. Gonçalves da Silva 4 (VIC). Viçosa, ESAV, 24-V-1935, J.G. Kuhlmann s.n. (VIC-1903); Viçosa, Campus da UFV, Recanto das Cigarras, 1-VI-1986, C.C. de Paula s.n. (VIC-9652); Viçosa, Campus da UFV, Silvicultura, 16-XII-1986, G.E. Valente & A.A. Azevedo 1 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, Estação Mata do Paraíso, 20º48’04”S, 42º51’19”W, 800 m, 23-V-2013, P.B. Schwartsburd & L. Rabelo-Sales 2799 (UCS, VIC); Viçosa, continuação da P.H. Rolfs, próximo ao condomínio novo, 25-IV-2016, P.B. Schwartsburd et al. 3776 (UCS, VIC); Viçosa, continuação da P.H. Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd et al. 3779 (UCS, VIC); Viçosa, continuação da P.H. Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd & N.S. Smith-Braga 3795 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, Estação Mata do Paraiso, Trilha dos Alpes, 9-II-2017, P.B. Schwartsburd. Et al. 3875 (VIC); Viçosa, Recanto das Cigarras, entre as churrasqueiras e as Bandeiras, 28-III-2019, V.S. Miranda et al. 55 (VIC); Viçosa, Recanto das Cigarras, entre as churrasqueiras e as Bandeiras, 28-III-2019, V.S. Miranda et al. 57 (VIC); Viçosa, Recanto das Cigarras, 28-III-2019, V.S. Miranda et al. 60 (VIC); Viçosa, Recanto das Cigarras, 28-III-2019, V.S. Miranda et al. 62 (VIC); Viçosa, estrada Viçosa-Cajuri, BR 121, Córrego Canela, a caminho do Sitio da EVA, 31-III-2019, V.S. Miranda et al. 72 (UCS, VIC).

Sticherus lanuginosus (Moric. Ex Feé) Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 20. 1950. Mertensia lanuginosa Moric. Ex Fée, Crypt. Vasc. Brésil 1. 202. 1869. Gleichenia lanuginosa (Moric. Ex Fée) Hieron., Hedwigia 48: 287. 1909.

Figures 2g-l

Plants terrestrial. Rhizomes brown, 2.4-5.5 mm diam; scales brown, lanceate, with dentate to serreate margins. Leaves erect; petioles 50-90 cm long, 3.1-4.5 mm diam, greenish to stramineous, with stramineous, fimbriate scales and arachnoid hairs; rachises sulcate, with stramineous or brown scales; buds with stramineous scales; ultimate segments two times longer than wide; midveins abaxially with fimbriate scales and arachnoid hairs, adaxially glabrous; veins 1-forked, abaxially with fimbriate scales and arachnoid hairs, adaxially glabrous; laminar tissue between the veins abaxially with fimbriate scales and arachnoid hairs, adaxially glabrous; laminar margins plane to revolute.

Distribution: Widespread in South America (and also in Costa Rica). In Viçosa it is found on road-sides, disturbed areas, forest edges and ravines, under full sunlight exposure.

Material examined: BRAZIL. Minas Gerais: Viçosa, 1930, Y. Mexia 4856 (F, MICH, NY, UC, US, VIC); Viçosa, ESAV, 24-II-1959, H.S. Irwin 2730 (MICH, NY, TEX, UC, US, VIC); Viçosa, Belvedere, 11-IX-1977, “Rosane & Rosângela” s.n. (VIC-7937); Viçosa, Campus da UFV, Silvicultura, 6-XII-1986, G.E. Valente & A.A. Azevedo 4 (VIC); Viçosa, Campus da UFV, Silvicultura, 23-II-1987, G.E. Valente & A.A. Azevedo 12 (VIC); Viçosa, Campus da UFV, Mata da Biologia, 10IX-1998, G.E. Valente & L.A. Basílio 353 (VIC); Viçosa, continuação da PH Rolfs, próximo ao condomínio novo, 25-IV-2016, P.B. Schwartsburd et al. 3775 (UCS, VIC); Viçosa, continuação da P.H. Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd et al. 3780 (UCS, VIC); Viçosa, continuação da P.H. Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd & N.S. Smith-Braga 3793 (UCS, VIC); Viçosa, continuação da P.H. Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd & N.S. Smith-Braga 3794 (VIC); Viçosa, Universidade Federal de Viçosa, estrada para criação de Equídeos, 13-X-2016, P.B. Schwartsburd & N.S. Smith-Braga 3799 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, Estação Mata do Paraíso, Trilha dos Alpes, 9-II-2017, P.B. Schwartsburd et al. 3876 (VIC); Viçosa, Universidade Federal de Viçosa, Recanto das Cigarras, 29-III-2019, V.S. Miranda et al. 64 (UCS, VIC); Viçosa, Universidade Federal de Viçosa, Recanto das Cigarras, 29-III-2019, V.S. Miranda et al. 65 (UCS, VIC); Viçosa, estrada Viçosa a Cajuri, BR 120, Córrego Canela, a caminho do Sitio da EVA, 31-III-2019, V.S. Miranda et al. 70 (VIC); Viçosa, estrada Viçosa a Cajuri, BR 120, em frente ao Hare Krishna, 31-III-2019, V.S. Miranda et al. 73 (VIC); Viçosa, Universidade Federal de Viçosa, 31-III-2019, V.S. Miranda et al. 78 (VIC).

Gonzales & Kessler (2011Gonzales, J. & Kessler, M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1-54.) and Lima & Salino (2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.) made some errors in the nomenclatural paragraph of S. lanuginosus. They cited the basinonym as “Gleichenia lanuginosa Moric. ex Fée”, but this name was never published by Fée. Instead, Fée (1869Fée, A.L.A. 1869. Cryptogames Vasculaires (fougères, lycopodiacées, hydroptéridées, équisétacées) du Brésil. J.B. Baillière et fils, Libraires, Paris.: 202) published Mertensia lanuginosa. The name Gleichenia lanuginosa was then combined by Hieronymus (1909Hieronymus, G. 1909. Plantae Stübelliana. Pteridophyta, vierter teil. Hedwigia 48: 215-303.), which was omitted by Gonzales & Kessler (2011Gonzales, J. & Kessler, M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1-54.) and Lima & Salino (2018Lima, L.V. & Salino, A. 2018. The fern family Gleicheniaceae (Polypodiopsida) in Brazil. Phytotaxa 358: 199-234.). Furthermore, these authors also cited “Gleichenia pennigera var. lanuginosa (Moric. Ex Fée) T. Moore” as a homotypic synonym of S. lanuginosus. In fact, Gleichenia pennigera var. lanuginosa T. Moore is a nomen nudum, which Moore (1862) was trying to publish as a new variety of Gleichenia pennigera, and in no way related to M. lanuginosa, which is a later name.

Sticherus lanuginosus is the most common Sticherus in southern and southeastern Brazil (pers. obs.), and is characterized by the erect leaves, and the ultimate segments about two times longer than wide. The other Sticherus from Viçosa have prostate or scandent leaves, and the ultimate segments are about four times longer than wide.

Sticherus nigropaleaceus (J.W. Sturm) J. Prado & Lellinger, Amer. Fern J. 86(3): 98. 1996. Mertensia nigropaleaceae J.W. Sturm, Fl. Bras. (Martius) 1(2): 222. 1859. Gleichenia bifida var. nigropaleacea (J.W. Sturm) Rosenst., Hedwigia 46: 60. 1906. Sticherus longipinnatus var. nigropaleaceus (J.W. Sturm) Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 22. 1950.

Figures 3a-f

Plants terrestrial. Rhizomes blackish, 2.5-3 mm diam; scales blackish, triangular, with dentate margins. Leaves scandent; petioles 40-72 cm long, 3.5-4 mm diam, blackish or reddish, glabrous; rachises with blackish scales with setose margins; buds with blackish scales; ultimate segments about four times longer than wide; midveins abaxially with stramineous scales and arachnoid hairs, adaxially glabrous; veins abaxially with stramineous scales and arachnoid hairs, adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; laminar margins plane to revolute.

Distribution: Endemic to Brazil, from Bahia to Rio Grande do Sul. In Viçosa it is found on road-sides, disturbed areas, forest edges and ravines, under full or part-time sunlight exposure.

Material examined: BRAZIL. Minas Gerais: Viçosa, Fazenda da Aguada, Buraco Frio, 725 m, 2-X-1930, Y. Mexia 5137 (F, MICH, NY, TEX, US, UC, VIC); Viçosa, Campus da UFV, Mata da Biologia, 10-IX-1998, G.E. Valente & L.A. Basílio 351 (VIC); Viçosa, Campus da UFV, final da Mata da Biologia, 20°45’31”S, 42°51’56”W, 16-VIII-2013, P.B. Schwartsburd et al. 2828 (UCS, VIC); Viçosa, continuação da PH Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd & N.S. Smith-Braga 3792 (VIC); Viçosa, continuação da PH Rolfs, próximo ao condomínio novo, 2-V-2016, P.B. Schwartsburd & N.S. Smith-Braga 3796 (VIC); Viçosa, nova P.H. Rolfs, no barranco do condomínio novo, 3-X-2019, P.B. Schwartsburd 4605 (UCS, VIC); Viçosa, UFV, Recanto das Cigarras, 20°48’30”S, 42°50’54”W, 29-III-2019, V.S. Miranda et al. 61 (VIC); Viçosa, UFV, Recanto das Cigarras, 20°48’30”S, 42°50’54”W, 29-III-2019, V.S. Miranda et al. 63 (UCS, VIC); Viçosa, UFV, Recanto das Cigarras, 20°48’30”S, 42°50’54”W, 29-III-2019, V.S. Miranda et al. 63b (VIC); Viçosa, UFV, Recanto das Cigarras, 20°48’30”S, 42°50’54”W, 29-III-2019, V.S. Miranda et al. 67 (VIC); Viçosa, UFV, Recanto das Cigarras, 20°48’30”S, 42°50’54”W, 29-III-2019, V.S. Miranda et al. 68 (VIC); Viçosa, estrada Viçosa a Cajuri, BR 121, Córrego Canela, a caminho do Sitio da EVA, 31-III-2019, V.S. Miranda et al. 71 (VIC).

Osmundales

Plants terrestrial or from paludose soils; corms with ectophloic siphonosteles, with a ring of discrete xylem strands; leaves dimorphic or hemidimorphic; sporangia large, with lateral annulli, opening by an apical split (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Smith et al. 2006Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.).

Osmundaceae Martinov

Leaves erect, dimorphic or hemi-dimorphic; laminae 1-2-pinnate; veins free; sporangia not forming sori, born along veins or covering the whole laminar abaxial surface; spores green, subglobose, trilete (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Smith et al. 2006Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2008. Fern classification. In: T.A. Ranker & C.H. Haufler (eds.). The biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge . pp. 417-467.; Sylvestre et al. 2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.).

Six genera: Claytosmunda (not occurring in Brazil), Leptopteris (not occurring in Brazil), Osmunda, Osmundastrum, Plenasium (not occurring in Brazil), and Todea (not occurring in Brazil).

Osmunda L.

Leaves hemi-dimorphic (with the fertile pinnae disposed on the distal 2/3 of the laminae) (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to tropical America. Springer-Verlag, New York.; Kramer & Green 1990Kramer, G. & Green, P. S. 1990. Pteridophytes and Gymnosperms. In: K. Kubitzki (ed.). The Families and Genera of Vascular Plants. Springer-Verlag, Vol. 1, Berlin.; Sylvestre et al. 2022Sylvestre, L.S., Costa, F.S.N. & Arana, M.D. 2022. A revision of the fern family Osmundaceae in Brazil. Acta Botanica Brasilica 36: e2020abb0537.).

Osmunda palustris Schrad., Gött. Gel. Anz.: 866. 1824. Osmunda spectabilis var. palustris (Schrad.) C. Presl, Supp. Tent. Pterid. 63. 1845. Osmunda regalis var. palustris (Schrad.) C. Chr. ex Angely, Fl. Paraná 23: 10. 1963. Osmunda regalis subsp. palustris (Schrad.) Á. Löve & D. Löve, Taxon 26(2, 3): 324. 1977.

Figures 4a-e

Corms erect, massive. Leaves hemi-dimorphic, fertile in the distal 1/3. Fertiles leaves 70-100(-120) cm long; petioles 20-40 cm × 3.2-4.5 mm, adaxially sulcate, glabrous; laminae 2-pinnate, 50-80 × 30-60 cm, chartaceous to coriaceous; rachises glabrous; pinnae in 7-12 pairs, opposite to sub-opposite, strongly ascending, imbricate, tappering at the apexes, producing apical pinnules 1/3 the length of the lateral pinnules; pinnules opposite to sub-opposite, articulate, petiolulate; petiolules with reddish hairs; costae abaxially and adaxially glabrous; veins abaxially strongly raised, abaxially and adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; laminar margins serrulate, glabrous.

Distribution: Likely widespread in the Neotropics. In Viçosa it is found on flooded soils inside forest or in forest edges, under shade or part-time sunlight exposure.

Material examined: BRAZIL. Minas Gerais: Piranga, entre Piranga e São João de Pirapitinga, pŕoximo ao Bar do Arlindo, 21-X-2017, P.B. Schwartsburd et al. 4369 (VIC); Viçosa, Fazenda da Aguada, 700 m, 14-X-1930, Y. Mexia 5169 (IAN, MO, NY, PO, U, UC, US, VIC); Viçosa, 14-I-1935, J.G. Kuhlmann s.n. (RB, VIC-1905); Viçosa, Mata da Prefeitura, 8-V-1978, M.P. Coons et al. 78-448 (VIC); Viçosa, Mata do Paraíso, 27-VI-2014, P.B. Schwartsburd et al. 3281 (UCS, VIC); Viçosa, Mata do Seu Nico, 15-XII-2014, P.B. Schwartsburd et al. 3456 (VIC).

Additional material examined: BRAZIL. Espírito Santo: Domingos Martins, 27-VI-2014, P.B. Schwartsburd et al. 3281 (VIC). Minas Gerais: Araponga, Parque Estadual da Serra do Brigadeiro, 1400 m, 5-V-2017, P.B. Schwartsburd et al. 4161 (UCS, VIC); Simonésia, RPPN Mata do Sossego, 19-IX-2014, P.B. Schwartsburd et al. 3350 (VIC). Paraná: Curitiba, Parque Municipal Tingui, 10-VII-2007, P.B. Schwartsburd 1354 (UPCB); Francisco Beltrão, 25-X-2006, P.B. Schwartsburd et al. 1134 (UPCB); Ponta Grossa, Parque Estadual de Vila Velha, 22-X-2004, P.B. Schwartsburd & R. Nagata 351 (HUPG, MBM, SP, UPCB).

Material examined of extra-Brazilian Osmunda: Osmunda regalis L., Sp. Pl. 2: 1065. 1753.

Material examined: ITALY. Piedmont: Torino, Giardino Reale di Savoia, 2-XII-2022, P.B. Schwartsburd 4903 (VIC); Torino, Giardino Reale di Savoia, 2-XII-2022, P.B. Schwartsburd 4905 (VIC).

Osmunda spectabilis Willd., Sp. Pl., ed. 4 [Willdenow] 5: 98. 1810. Osmunda regalis var. spectabilis (Willd.) A. Gray, Manual (Gray), ed. 2. 600. 1856. Osmunda regalis subsp. spectabilis (Willd.) Á. Löve & D. Löve, Taxon 26(2, 3): 324. 1977.

Figures 4f-g

Material examined: UNITED STATES OF AMERICA. Vermont: outside Burlington, 10-IX-2018, P.B. Schwartsburd et al. 4471 (VIC, VT); Waterbury, around the Reservoir, 10-VII-2018, P.B. Schwartsburd & M.P. Schwartsburd 4449 (VIC, VT).

Acknowledgments

We thank Andreza Oliveira, Cecília Miranda, Nayara Smith-Braga, Nelson Túlio Pena, and Tatiane Campos, for helping with field work. Pedro Schwartsburd also thanks Art Guillman, Jakob Suissa, Mauro Schwartsburd, Miodeli Nogueira Jr., Renato Nagata, and Susan Fawcett for helping with field work, and the staff of the Pringle Herbarium (VT), for handling and shipping the collections from Vermont, especially Eunice Froeliger.

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