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Fern flora of Viçosa, Minas Gerais State, Brazil: Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae

Flora de samambaias de Viçosa, Estado de Minas Gerais, Brasil: Dennstaedtiaceae, Lindsaeaceae e Saccolomataceae

ABSTRACT

As part of an ongoing project treating the ferns and lycophytes from the region of Viçosa, Minas Gerais State, Brazil, we here present the taxonomic treatment of the early divergent lineages of the leptosporangiate ferns: the families Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae. We have been sampling the remnant forest patches since 2012; we also fully reviewed the collection of herbarium VIC and other online collections: F, IAN, NY, PH, RB, U, UC, UPCB, US, and WTU. In the region of Viçosa, six taxa belonging to those families occur: Dennstaedtia cicutaria and Pteridium esculentum subsp. arachnoideum (Dennstaedtiaceae); Lindsaea lancea var. lancea, L. quadrangularis subsp. quadrangularis, and L. stricta var. stricta (Lindsaeaceae); and Saccoloma elegans (Saccolomataceae). Among these taxa, only L. quadrangularis subsp. quadrangularis and S. elegans are endemic to the Brazilian Atlantic Rainforest; the remaining are widespread in the Neotropics. We present keys, descriptions, illustrations, examined specimens, and comments.

Keywords:
Brazilian Atlantic Rainforest; Polypodiales; pteridophytes; southeast Brazil; taxonomy

RESUMO

Como parte de um projeto em andamento sobre samambaias e licófitas da região de Viçosa, Estado de Minas Gerais, Brasil, apresenta-se o tratamento taxonômico das primeiras linhagens divergentes de samambaias leptosporangiadas: as famílias Dennstaedtiaceae, Lindsaeaceae e Saccolomataceae. Expedições de campo foram realizadas desde 2012 em remanescentes florestais da região. Também foi realizada uma revisão completa da coleção do herbário VIC, bem como de outras coleções online de herbários: F, IAN, NY, PH, RB, U, UC, UPCB, US, and WTU. Na região de Viçosa, existem seis táxons pertencentes a essas famílias: Dennstaedtia cicutaria e Pteridium esculentum subsp. arachnoideum (Dennstaedtiaceae); Lindsaea lancea var. lancea, L. quadrangularis subsp. quadrangularis e L. stricta var. stricta (Lindsaeaceae); e Saccoloma elegans (Saccolomataceae). Dentre esses táxons, apenas L. quadrangularis subsp. quadrangularis e S. elegans são endêmicos da Mata Atlântica brasileira; as demais estão amplamente distribuídas na região Neotropical. São apresentadas chaves, descrições, ilustrações, materiais examinados e comentários.

Palavras-chave:
Mata Atlântica brasileira; Polypodiales; pteridófitas; sudeste do Brasil; taxonomia

Introduction

At the end of the era in which the plant Classification Systems were mostly based on morphological data, Dennstaedtiaceae was considered a big family, comprehending three subfamilies (or tribes) and about 17 genera (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to Tropical America. Springer-Verlag, New York., Kramer 1990Kramer, K.U. 1990. Dennstaedtiaceae. In: K.U. Kramer, P.S. Green (eds.), v. I, Pteridophytes and Gymnosperms, pp. 81-94. In: K. Kubitzki (ed.). The families and genera of vascular plants. Springer-Verlag, Berlin, Germany.). With the advent of molecular data, those subfamilies were raised to family-level, some families were created, and now the early-diverging leptosporangiate ferns are represented by Cystodiaceae (one genus), Dennstaedtiaceae with a stricter circumscription (11-12 genera), Lindsaeaceae (seven genera), Lonchitidaceae (one genus), Saccolomataceae (one or two genera), as well as the great Pteridaceae (+50 genera) (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731., PPG I 2016PPG I - Pteridophyte Phylogeny Group. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603., Shang et al. 2018Shang, H., Sundue, M., Wei, R., Wei, X.P., Luo, J.J., Liu, L., Schwartsburd, P.B., Yan, Y.H. & Zhang, X.C. 2018. Hiya: A new genus segregated from Hypolepis in the fern family Dennstaedtiaceae, based on phylogenetic evidence and character evolution. Molecular Phylogenetics and Evolution 127: 449-458., Schwartsburd et al. 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. ).

Brazil is represented by Dennstaedtiaceae with eight genera and 26 species, Lindsaeaceae with one genus and 34 spp., Lonchitidaceae with one genus and one sp., and Saccolomataceae with one genus and nine spp., leaving Pteridaceae apart (which is subject to another paper) (Kramer 1957Kramer, K.U. 1957. A revision of the genus Lindsaea in the new world with notes on allied genera. Acta Botanica Neerlandica 6: 97-290., Pena et al. 2020Pena, N.T.L., Miranda, C.V., Schwartsburd, P.B., Oliveira, A.G.S. & Smith-Braga, N. 2020. Lindsaeaceae in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available in http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91283 (access in 24-XII-2021).
http://floradobrasil.jbrj.gov.br/reflora...
, Schwartsburd 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. , Schwartsburd & Pena 2020Schwartsburd, P.B. & Pena, N.T.L. 2020. Dennstaedtiaceae in Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. Available in https://floradobrasil.jbrj.gov.br/FB90915 (access in 26-VIII-2022).
https://floradobrasil.jbrj.gov.br/FB9091...
). In general, these plants present characters considered plesiomorphic that are typical of the early-diverging lineages of Polypodiales, such as solesnotelic rhizomes clothed with hairs or primitive scales, large, highly dissected leaves, the sori that are truly marginal and protected by inner and outer indusia, and trilete spores (Tryon & Tryon 1982Tryon, R.M. & Tryon, A.F. 1982. Ferns and allied plants, with special reference to Tropical America. Springer-Verlag, New York., Kramer 1990Thomson, J.A. 2012. Taxonomic status of diploid southern hemisphere brackens (Pteridium: Dennstaedtiaceae). Telopea 14: 43-48., PPG I 2016PPG I - Pteridophyte Phylogeny Group. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603., Schwartsburd et al. 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. ).

Pteridium is spread all across the globe (except for Antarctic) and may be the one most important to mankind by if directly affects the local human populations: it behaves as a pioneer weed invading croplands; it spreads its underground rhizomes making it hard to be removed; it intoxicates the cattle and horses; and it also intoxicates humans, when ingested - some studies also pointed out the transmission of its toxins (the psilatoquinis) via cow milk (e.g., Tryon 1941Tryon, R.M. 1941. Revision of the genus Pteridium. Contributions from the Gray Herbarium of Harvard University 134 (reprint from Rhodora 43): 1-31, 37-67., Evans 1976Evans, I.A. 1976. Relationship between bracken and cancer. Botanical Journal of the Linnean Society 73: 105-112., Page 1976Page, C.N. 1976. The taxonomy and phytogeography of bracken. Botanical Journal of the Linnean Society 73: 1-34. , Marrs et al. 2000Marrs, R.H., Le Duc, M.G., Mitchell, R.J., Goddard, D., Paterson, S. & Pakerman, R.J. 2000. The ecology of bracken: Its role in succession and implications for control. Annals of Botany 85: 3-15., Martini-Santos et al. 2010Martini-Santos, B.J., Peixoto, T.C., Galvão, A., Prado, J.S., Aragão, A.P. & Benevides, M.L.S. 2010. Surto de intoxicação aguda por Pteridium arachnoideum (Kaulf.) em bovinos (Outbrake of acute poisoning by Pteridium arachnoideum (Kaulf.) in cattle). Revista Brasileira de Medicina Veterinária 32: 92-96., Oliveira et al. 2018Oliveira, V.M., Schwartsburd, P.B., Brighenti, A.M., d’Oliveira, P.S. & Miranda, J.E.C. 2018. Plantas tóxicas em pastagens: samambaia-do-campo (Pteridium esculentum subsp. arachnoideum (Kaulf.) Thomson, família Dennstaedtiaceae). Embrapa, Comunicado Técnico 84, Juiz de Fora., Schwartsburd et al. 2018Schwarstsburd, P.B, Yañez, A. & Prado, J. 2018. Formal recognition of six subordinate taxa within the South American bracken fern, Pteridium esculentum (P. esculentum subsp. arachnoideum s.l. - Dennstaedtiaceae), based on morphology and geography. Phytotaxa 333: 22-40.).

In the State of Minas Gerais, Dennstaedtiaceae is represented by 14 species, Lindsaeaceae by 11 spp., and Saccolomataceae by three spp. (Pena et al. 2020Pena, N.T.L., Miranda, C.V., Schwartsburd, P.B., Oliveira, A.G.S. & Smith-Braga, N. 2020. Lindsaeaceae in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available in http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91283 (access in 24-XII-2021).
http://floradobrasil.jbrj.gov.br/reflora...
, Schwartsburd 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. , Schwartsburd & Pena 2020Schwartsburd, P.B. & Pena, N.T.L. 2020. Dennstaedtiaceae in Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. Available in https://floradobrasil.jbrj.gov.br/FB90915 (access in 26-VIII-2022).
https://floradobrasil.jbrj.gov.br/FB9091...
, Schwartsburd 2022Schwartsburd, P.B. 2022. Saccoloma (Saccolomataceae) from Brazil, including new species and new combinations. Brittonia 74: 155-179.). Here, we present the taxonomic treatment of these families for the region of Viçosa, Minas Gerais State, Brazil, as part of the ongoing project on the Fern Flora of this region with fascicles in preparation and others already published: Miranda & Schwartsburd (2016Miranda, C.V. & Schwartsburd, P.B. 2016. Aquatic ferns from Viçosa (MG, Brazil): Salviniales (Filicopsida; Tracheophyta). Brazilian Journal of Botany 39: 935-942. - Salviniales), Rabelo & Schwartsburd (2016Rabelo, L.S. & Schwartsburd, P.B. 2016. Schizaeales (Filicopsida, Tracheophyta) of Viçosa, Minas Gerais, Brazil, with special reference to hybrids. Brittonia 68: 379-396. - Schizaeales), and 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. - Polypodiaceae).

Material and methods

In the region of Viçosa, Minas Gerais State, Brazil, the remaining forest patches are classified as Semi-Deciduous Seasonal Forest (IBGE 2012IBGE - Instituto Brasileiro de Geografa e Estatística. 2012. Manual técnico da vegetação Brasileira. Série Manuais Técnicos em Geociências , 2ª ed. IBGE, Rio de Janeiro.). The local elevation ranges from 600 to 900 m.a.s.l. (pers. obs.), and the annual temperatures range from 5° to 30°C (INMET 2021INMET - Instituto Nacional de Meteorologia. 2021 - Continuous update. Ministério da Agricultura, Pecuária e Abastecimento. Available in http://www.inmet.gov.br/portal/ (access in 29-VIII-2021).
http://www.inmet.gov.br/portal/...
).

We have been sampling the remnant forest patches since 2012. Specimens were collected and dried according to usual methods for fern collections (Windisch 1992Windisch, P.G. 1992. Pteridófitas da região norte-ocidental do Estado de São Paulo: guia para estudo e excursões. 2 ed. Universidade Estadual Paulista, São José do Rio Preto.) and then incorporated in herbarium VIC (acronyms following Thiers, continuously updatedThiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available in http://sweetgum.nybg.org/science/ih/ (access in 26-VIII-2022).
http://sweetgum.nybg.org/science/ih/ ...
), with duplicates to be sent to other herbaria. We also fully reviewed the collection of herbarium VIC and collections of other herbaria online: F, IAN, NY, PH, RB, U, UC, UPCB, US, and WTU. Although the classification system adopted for the previous fascicles was based on an adaptation of Kenrick & Crane (1997Kenrick, P. & Crane, P.R. 1997. The origin and early diversification of land plants. A cladistic study. Smithsonian Institution Press, Washington D.C., USA.) and Smith et al. (2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.), we here adopt the system of PPG I (2016PPG I - Pteridophyte Phylogeny Group. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603.). Morphological terms are according to Lellinger (2002Lellinger, D.B. 2002. A modern multilingual glossary for taxonomic Pteridology. Pteridologia 3: 1-263. ).

Results and Discussion

Dennstaedtiaceae, Lindsaeaceae, and Saccolomataceae, presented a total of four genera and six species occurring in the region of Viçosa. Four of those species have also infra-specific classifications, which were adopted here. The six taxa are: Dennstaedtia cicutaria (Sw.) T. Moore, and Pteridium esculentum subsp. arachnoideum (Kaulf.) J.A. Thomson (Dennstaedtiaceae); Lindsaea lancea (L.) Bedd. var. lancea, L. quadrangularis Raddi subsp. quadrangularis, and L. stricta (Sw.) Dryand. var. stricta (Lindsaeaceae); and Saccoloma elegans Kaulf. Among them, only L. quadrangularis subsp. quadrangularis and S. elegans are endemic to the Brazilian Atlantic Rainforest; the remaining taxa are widespread in the Neotropics. Differently from the other studied families: Salviniaceae (Miranda & Schwartsburd 2016Miranda, C.V. & Schwartsburd, P.B. 2016. Aquatic ferns from Viçosa (MG, Brazil): Salviniales (Filicopsida; Tracheophyta). Brazilian Journal of Botany 39: 935-942.); Anemiaceae (Rabelo & Schwartsburd 2016Rabelo, L.S. & Schwartsburd, P.B. 2016. Schizaeales (Filicopsida, Tracheophyta) of Viçosa, Minas Gerais, Brazil, with special reference to hybrids. Brittonia 68: 379-396. ); Polypodicaeae (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 have not recollected half of the taxa involved in this paper, which were collected on the 1930s and never again. Probably, they went locally extinct due to forest fragmentation and disturbance. We have not recollected D. cicutaria, L. lancea var. lancea, nor L. quandrangularis subsp. quadrangularis.

  • Key to taxa of Dennstaedtiaceae, Lindsaeaceae and Saccolomataceae from the region of Viçosa

  • 1. Sori born at the tip of single veins

  • 2. Laminae 1-pinnate; sori along the laminar margins, parallel to each other ………………………………………………………….…. Saccoloma elegans (Saccolomataceae)

  • 2. Laminae 3-pinnate-pinnatifid to 4-pinnate; sori born on sinuses of the segments …………………………………………………...……. Dennstaedtia cicutaria (Dennstaedtiaceae)

  • 1. Sori born at the tip of multiple veins (on a commisural vein)

  • 3. Rhizomes with hairs; laminae 3-4-pinnate-pinnatifid …………………………………………. …………………………………. Pteridium esculentum subsp. arachnoideum (Dennstaedtiaceae)

  • 3. Rhizomes with scales; laminae 1-2-pinnate (or rarely 3-pinnate) ………….……. (Lindsaeaceae)

  • 4. Stipes abaxially terete, adaxially flattened ………………….……. Lindsaea stricta var. stricta

  • 4. Stipes quadrangular in cross section

  • 5. Stipes and rachises reddish-brown; sori interrupted …………………………… ………………………………...……...…... Lindsaea quandrangularis subsp. quadrangularis

  • 5. Stipes proximally dark brown, stramineous above; rachises stramineous; sori not interrupted …………………………………………………..….……………. Lindsaea lancea var. lancea

Dennstaedtiaceae

Rhizomes generaly long-creeping, solenostelic or rarely dictyostelic or polycyclic, bearing hairs or rarely with primitive scales. Leaves monomorphic; stipes often with epipetiolar buds, usually with a omega-shaped vascular strand; laminae 2-5-pinnate, rarely less dissected; veins free or rarely anastomosing and then without included veinlets; indument formed by hairs or glandular hairs, or absent. Sori marginal or submarginal, linear or discrete, introrse or extrorse, protected by outer and/or inner indusia or rarely not protected; spores tetrahedral and trilete, or reniform and monolete (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.).

The family is cosmopolitan, composed of eleven genera and ca. 270 species (PPG I 2016PPG I - Pteridophyte Phylogeny Group. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603., Shang et al. 2018Shang, H., Sundue, M., Wei, R., Wei, X.P., Luo, J.J., Liu, L., Schwartsburd, P.B., Yan, Y.H. & Zhang, X.C. 2018. Hiya: A new genus segregated from Hypolepis in the fern family Dennstaedtiaceae, based on phylogenetic evidence and character evolution. Molecular Phylogenetics and Evolution 127: 449-458., Schwartsburd et al. 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. ). The clade containing Dennstaedtia, Leptolepia, Microlepia, and Oenotrichia still needs recircumscriptions, due to a polyphyletic Dennstaedtia (Schwartsburd et al. 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. ).

Dennstaedtia Bernh.

Rhizomes short to long-creeping, glabrous or with catenate hairs. Leaves large, to 4 m long; laminae 2-4-pinnate; veins free; indument formed by catenate hairs, acicular hairs, or absent; sori marginal, discrete, at the tip of single veins; inner and outer indusia fused into purse- or cup-shaped or hemi-globose indusia (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.).

Dennstaedtia is polyphyletic in current circumscription (Schwartsburd et al. 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. ), with about 70 species. In Brazil occur six species (Schwartsburd & Pena 2020Pena, N.T.L., Miranda, C.V., Schwartsburd, P.B., Oliveira, A.G.S. & Smith-Braga, N. 2020. Lindsaeaceae in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available in http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91283 (access in 24-XII-2021).
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), but there are probably about eight to ten species (pers. obs.).

Dennstaedtia cicutaria (Sw.) T. Moore, Index Fil. (T. Moore) 2: xcvii. 1857.

Figure 1a-c

Figure 1. a-c.
Dennstaedtia cicutaria (Sw.) T. Morre. a. rhizome. b. basal pinna. c. segments, abaxially, showing indument and sori. d-g. Pteridium esculentum subsp. arachnoideum (Kaulf.) J.A. Thomson. d. rhizome. e. pinnule. f. segment, abaxially, showing indument and sori. g. segment in cross section, showing indument and outer indusium (abaxial surface up).

Plants terrestrial. Rhizomes long-creeping, with brownish catenate hairs. Leaves 1-3.5 m long; stipes 0.5-2.5 m long, proximally brown, lighter brown above, with acicular hairs or glabrescent; laminae 0.5-2 m long, proximally 3-pinnate-pinnatifid to 4-pinnate; rachises light brown, pilose, with brownish catenate hairs, without proliferous buds; basal pinnae 25-100 cm × 12-35 cm, subequilateral; pinnules sessile, lanceolate to oblong, margins dentate; costae abaxially and adaxially pilose, with light brown to reddish catenate hairs; costules abaxially and adaxially pilose, with light brown to reddish catenate hairs; veins free, abaxially and adaxially hirsute, with erect, light brown to reddish acicular hairs; laminar tissue between the veins abaxially and adaxially with catenate and acicular hairs; sori oblong or rounded, born at the tip of single veins, in sinuses of segments; indusia purse-shaped, commonly with acicular and catenate hairs.

Geographical distribution: Neotropical.

Specimens examined: BRAZIL. Minas Gerais: Viçosa: Agricultural College land, Horticultural Hill, 670 m, 7-VII-1930, Y. Mexia 4846 (F, MO-n.v., U, UC, US); Escola Superior de Agricultura e Veterinária, base of Horticultural Hill, 668 m, 17-VII-1930, Y. Mexia 4866 (IAN, F, U, UC, US, VIC, WTU); Escola Superior de Agricultura e Veterinária, I-1934, J.G. Kuhlmann s.n. (VIC-1309).

Dennstaedtia cicutaria is recognized by veins and laminar tissue between the veins densely with catenate and acicular hairs, indusia purse-shaped with acicular and catenate hairs.

Pteridium Gled. ex Scop., nom. cons.

Rhizomes long-creeping, with setose hairs. Leaves large, to 4.5 m long; laminae 3-5-pinnate; veins free; indument formed by catenate hairs, acicular hairs, arachnoid hairs, and/or minute gnarled hairs; sori marginal, linear, continuous along the laminar margins, born on comissural veins; inner indusia vestigial; outer indusia linear (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.).

Two species plus two species with hybrid origin are currently recognized (Thomson 2012Thomson, J.A. 2012. Taxonomic status of diploid southern hemisphere brackens (Pteridium: Dennstaedtiaceae). Telopea 14: 43-48., Zhou et al. 2014 Zhou, S., Dong, W., Chen, X., Zhang, X., Wen, J. & Schneider, H. 2014. How many species of bracken (Pteridium) are there? Assessing the Chinese brackens using molecular evidence. Taxon 63: 509-521., Schwartsburd et al. 2018Schwarstsburd, P.B, Yañez, A. & Prado, J. 2018. Formal recognition of six subordinate taxa within the South American bracken fern, Pteridium esculentum (P. esculentum subsp. arachnoideum s.l. - Dennstaedtiaceae), based on morphology and geography. Phytotaxa 333: 22-40., 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. ). In addition, a total of ca. 20 morpho-taxa are also recognized as infra-specific entities, such as subspecies, varieties, forms, or combinations of more than one rank (e.g., Tryon 1941Tryon, R.M. 1941. Revision of the genus Pteridium. Contributions from the Gray Herbarium of Harvard University 134 (reprint from Rhodora 43): 1-31, 37-67., Page 1976Page, C.N. 1976. The taxonomy and phytogeography of bracken. Botanical Journal of the Linnean Society 73: 1-34. , Thomson 2012Thomson, J.A. 2012. Taxonomic status of diploid southern hemisphere brackens (Pteridium: Dennstaedtiaceae). Telopea 14: 43-48., Schwartsburd et al. 2018Schwarstsburd, P.B, Yañez, A. & Prado, J. 2018. Formal recognition of six subordinate taxa within the South American bracken fern, Pteridium esculentum (P. esculentum subsp. arachnoideum s.l. - Dennstaedtiaceae), based on morphology and geography. Phytotaxa 333: 22-40.). In Brazil, the following taxa are currently recognized: Pteridium caudatum (L.) Maxon, P. esculentum subsp. arachnoideum, P. esculentum subsp. campestre (Schrad.) Schwartsb. & J. Prado, P. esculentum subsp. gryphus Schwartsb., P. esculentum var. harpianum Schwartsb. & A. Yañez, and P. esculentum var. paedomorficum Schwartsb. & J. Prado (Schwartsburd 2020Schwartsburd, P.B. 2020. Saccolomataceae in Flora e Funga do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available in http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB92021 (access in 24-XIII-2021).
http://floradobrasil.jbrj.gov.br/reflora...
). The most common taxon in southeastern Brazil is P. esculentum subsp. arachnoideum, which invades crops and is toxic for cattle, horses, and human ingestion (Schwartsburd et al. 2018Schwarstsburd, P.B, Yañez, A. & Prado, J. 2018. Formal recognition of six subordinate taxa within the South American bracken fern, Pteridium esculentum (P. esculentum subsp. arachnoideum s.l. - Dennstaedtiaceae), based on morphology and geography. Phytotaxa 333: 22-40.).

Pteridium esculentum subsp. arachnoideum (Kaulf.) J.A. Thomson, Telopea 14: 45. 2012Thomson, J.A. 2012. Taxonomic status of diploid southern hemisphere brackens (Pteridium: Dennstaedtiaceae). Telopea 14: 43-48..

Figure 1 d-g

Plants terrestrial, ticket-forming. Rhizomes long-creeping, with ciliform hairs. Leaves 1.5-4.5m long; stipes 0.75-2.5 m long, proximally dark brown, stramineous above, glabrous; laminae 0.75-2.5m m long, proximally 3-4-pinnate-pinnatifid; rachises stramineous, glabrous; basal pinnae 60-100 × 25-70cm, equilateral; pinnae and pinnules distally with free lobes between the segments; compound distal segments inequilateral, irregularly dissected, caudate at apex; simple distal segments linear, up to 3(-4) cm long; costae abaxially with reddish catenate hairs, adaxially glabrous; costules abaxially with reddish catenate hairs, adaxially glabrous; veins abaxially lanose, with lax, arachnoid hairs, adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; sori marginal, linear, along segment margins, born at the tip of multiple veins; inner indusia vestigial; outer indusia linear.

Geographical distribution: Neotropical.

Specimens examined: BRAZIL. Minas Gerais: Viçosa: Entre Viçosa e Visconde do Rio Branco, 11-V-2013, P.B. Schwartsburd et al. 2837 (VIC); idem, 11-V-2013, P.B. Schwartsburd et al. 2838 (VIC); State Agricultural School, 24-II-1957, H.S. Irwin 2721 (F, TEX-n.v., UC, US, VIC); Escola Superior de Agricultura e Veterinária, 1943, P. Alvin s.n. (VIC-3537); Universidade Federal de Viçosa, Fazendinha, 11-IX-1977, Rosane & Rosângela s.n. (VIC-7962, 7963, 7964, 7965); idem, Mata do Paraíso, Trilha dos Alpes, 9-II-2017, P.B. Schwartsburd et al. 3877 (VIC); idem, Mata do Paraíso, Trilha dos Alpes, 9-II-2017, P.B. Schwartsburd et al. 3879 (VIC); idem, Mata da Biologia, 27-III-2019, N.T.L. Pena et al. 783 (VIC); idem, Mata da Biologia, 27-III-2019, N.T.L. Pena et al. 784 (VIC).

Pteridium esculentum subsp. arachnoideum is recognized by the pinnae and pinnules distally with free lobes between the segments, veins abaxially with arachnoid hairs, and glabrous laminar tissue between the veins. The outer indusia is always present, even in sterile leaves.

Lindsaeaceae

Rhizomes short to long-creeping, protostelic with internal phloem, or rarely solenostelic, bearing basally attached, nonclathrate scales or hairs. Leaves monomorphic; laminae simple to 3-pinnate or rarely more dissected; veins usually free, occasionally anastomosing and then without included veinlets; indument generally absent or formed by 2-celled glandular hairs. Sori marginal or submarginal, extrorse, elongate to linear, protected by outer (not modified) and inner (modified) indusia; spores tetrahedral and trilete, or rarely bilateral and monolete (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.).

Lindsaea Dryand. ex Sm.

Rhizomes short-creeping, with primitive scales. Leaves small to medium-sized, ca. 20 cm to 1 m long; laminae simple to 3-pinnate; indument absent or formed by 2-celled glandular hairs; veins generally free; sori marginal or submarginal, elongate, born on comissural veins; inner indusia modified, elongate; outer indusia not modified (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.).

About 180 species, with 34 spp. occurring in Brazil (Kramer 1957Kramer, K.U. 1957. A revision of the genus Lindsaea in the new world with notes on allied genera. Acta Botanica Neerlandica 6: 97-290., PPG I 2016PPG I - Pteridophyte Phylogeny Group. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603., Pena et al. 2020Pena, N.T.L., Miranda, C.V., Schwartsburd, P.B., Oliveira, A.G.S. & Smith-Braga, N. 2020. Lindsaeaceae in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available in http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB91283 (access in 24-XII-2021).
http://floradobrasil.jbrj.gov.br/reflora...
).

Lindsaea lancea (L.) Bedd. var. lancea, Suppl. Ferns S. Ind. 6.: 292. 1876.

Figure 2 a-d

Figure 2. a-d.
Lindsaea lancea (L.) Bedd. var. lancea. a. rhizome. b. petiole, in cross section. c. leaf. d. pinnule, abaxially, showing sorus. e-h. Lindsaea quadrangularis Raddi subsp. quadrangularis. e. rhizome. f. petiole, in cross section. g. leaf. h. pinnule, abaxially, showing sorus.

Plants terrestrial. Rhizomes short-creeping, with dark brown, lanceolate scales. Leaves 30-50 cm long; stipes 15-25 cm long, dark brown proximally, stramineous above, quadrangular, proximally with dark brown, acuminate scales and sparse 2-celled glandular hairs; laminae 15-25 cm long, 2-pinnate; rachises not winged, stramineous, with sparse 2-celled glandular hairs; basal pinnae 10-15cm × 2.5-3.5cm equilateral; pinnules petiolulate, dimidiate, 1.5-2 cm long; costae abaxially and adaxially with sparse 2-celled glandular hairs or glabrescent; veins abaxially and adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; sori continuous, born at the tip of multiple veins; inner indusia membranaceous to papyraceous, the margins entire, wavy, glabrous.

Geographical distribution: Neotropical.

Specimens examined: BRAZIL. Minas Gerais: Viçosa, 1930, Y. Mexia 4639-a (GH-n.v., UC, US, VIC).

All Lindsaea spp. from Viçosa are similar in having 2-pinnate laminae and dimidiate pinnules. Lindsaea lancea var. lancea differs from the other two by having continuous sori; L. quadrangularis subsp. quadrangularis differs from the other two by having reddish-brown, winged rachises; and L. stricta var. stricta differs from the other two by having petioles that are terete abaxially. In addition, pinnule lenghts are different between the three taxa: 1.5-2 cm in L. lancea var. lancea, 1.2-1.5 cm in L. quadrangularis subsp. quadrangularis, and to 1 cm in L. stricta var. stricta.

Lindsaea quadrangularis Raddi subsp. quadrangularis, Opusc. Sci. 3: 294. 1825.

Figure 2 e-h

Plants terrestrial. Rhizomes short-creeping, with light brown, lanceolate scales. Leaves 40-75 cm long; stipes 25-40 cm long, entirely reddish-brown, quadrangular, proximally with light brown, acuminate scales and sparse 2-celled glandular hairs; laminae 20-40 cm long, 2-pinnate; rachises winged, reddish-brown, with sparse 2-celled glandular hairs; basal pinnae 10-18 × 2-3 cm; pinnules short-petiolulate, dimidiate, 1.2-1.5 cm long; costae abaxially and adaxially with sparse 2-celled glandular hairs; veins abaxially and adaxially glabrous; laminar tissue between the veins abaxially with sparse 2-celled glandular hairs, adaxially glabrous; sori interrupted, born at the tip of multiple veins; indusia membranaceous, the margin dentate, glabrous.

Geographical distribution: endemic to the Brazilian Atlantic Forest.

Specimens examined: BRAZIL. Minas Gerais: Viçosa: Agricultural College lands, Hillside of Cha-Cha Valley, 1-III-1930, Y. Mexia 4410-a (UC, US-n.v.); s.l., 1935, J.G. Kuhlmann s.n. (VIC-1894).

Lindsaea stricta (Sw.) Dryand. var. stricta, Trans. Linn. Soc. London 3: 42. 1797.

Figure 3 a-d

Figure 3. a-d.
Lindsaea stricta (Sw.) Dryand. var. stricta. a. rhizome. b. petiole, in cross section. c. leaf. d. pinnule, abaxially, showing sorus. e-l. Saccoloma elegans Kaulf. e. rhizome. f. and g. petiolar scales. h. medium pinna. i. base of pinna. j. pinna, abaxially, showing venation pattern. k. pinna, abaxially, showing sori. l. apex of pinna.

Plants terrestrial. Rhizomes short-creeping, with dark brown, lanceolate scales. Leaves 35-80 cm long; stipes 15-50 cm long, dark brown proximally, stramineous above, abaxially terete, adaxially flattened, proximally with dark brown, acuminate scales; laminae 20-35cm long, 2-pinnate or rarely 1- or 3-pinnate; rachises not winged, stramineous, with sparse 2-celled glandular hairs or glabrescent; basal pinnae 12-25 cm × 1-2 cm; pinnules short-petiolate, dimidiate or flabellate, to 1 cm long; costae abaxially and adaxially with sparse 2-celled glandular hairs or glabrescent; veins abaxially and adaxially with sparse 2-celled glandular hairs or glabrous; laminar tissue between the veins abaxially and adaxially with sparse 2-celled glandular hairs or glabrous; sori interrupted, born at the tip of multiple veins; inner indusia membranaceous, the margins dentate, glabrous.

Geographical distribution: Neotropical.

Specimens examined: BRAZIL. Minas Gerais: Viçosa: road to Cajurí, Moro Grande, 750 m, 7-I-1931, Y. Mexia 5486 (B-n.v., BM-n.v., C-n.v., F, G-n.v., GH-n.v., K-n.v., MO-n.v., NY-n.v., PH-n.v., S-n.v., TEX-n.v., U-n.v., UC, US, VIC); Silvestre, 10-IV-1999, M.G. Bovini 1624 (VIC); Universidade Federal de Viçosa, Mata da Silvicultura, em direção ao Cristo, 740 m, 20º46’S, 42º52’W, 31-X-2014, P.B. Schwartsburd et al. 3421, 3423 (VIC); idem, Mata do Paraíso, Trilha dos Alpes, 9-II-2017, P.B. Schwartsburd et al. 3881 (VIC); idem, Recanto das Cigarras, 8-VII-2019, N.T.L. Pena et al. 1049 (VIC).

Saccolomataceae

Rhizomes erect or rarely short-creeping, sometimes forming small trunks, dictyostelic, bearing peltate scales. Leaves monomorphic; stipes with an omega-shaped vascular strand; laminae 1-4-pinnate; veins free; indument formed by caduceus scales or rarely by scattered catenate hairs. Sori marginal or submarginal, discrete, extrorse, protected by outer (not modified) and inner (modified) indusia; spores tetrahedral-globose and trilete (Smith et al. 2006Smith, A.R., Pryer, K.M., Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A Classification for Extant Ferns. Taxon 55: 705-731.).

One pantropical genus with about 24 species (Rojas-Alvarado 2010Rojas-Alvarado, A.F. 2010. Novelties in the Saccoloma inaequale complex (Saccolomataceae) from the Neotropics. Métodos en Ecología y Sistemática 5: 1-16., Luong et al. 2015Luong, T.T., Hovenkamp, P.H. & Sosef, M.S.M. 2015. Revision of the fern genus Orthiopteris (Saccolomataceae) in Malesia and adjacent regions. PhytoKeys 53: 39-71., Schwartsburd 2020Schwartsburd, P.B., Perrie, L.R., Brownsey, P., Shepherd, L.D., Shang, H., Barrington, D.S. & Sundue, M.A. 2020. New insights into the evolution of the fern family Dennstaedtiaceae from an expanded molecular phylogeny and morphological analysis. Molecular Phylogenetics and Evolution 150: 106881. , Schwartsburd 2022Schwartsburd, P.B. 2022. Saccoloma (Saccolomataceae) from Brazil, including new species and new combinations. Brittonia 74: 155-179.). The adoption of genus Orthiopteris for the Paleotropical species is phylogenetically justified, but lacks morphological support; thus, regarding only Saccoloma in Saccolomataceae is more convenient (Schwartsburd 2022Schwartsburd, P.B. 2022. Saccoloma (Saccolomataceae) from Brazil, including new species and new combinations. Brittonia 74: 155-179.). In Brazil, nine species are currently recognized (Schwartsburd 2022Schwartsburd, P.B. 2022. Saccoloma (Saccolomataceae) from Brazil, including new species and new combinations. Brittonia 74: 155-179.).

Saccoloma Kaulf.

Saccoloma elegans Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 51. 1820.

Figure 3 e-l

Plants terrestrial. Rhizomes erect to decumbent, stout to trunk-like, glabrescent; leaves 1.2-2.2 m long; petioles proximally burgundy, stramineous above, 40-80 cm long, proximally wrinkled to minutely spiny; petiolar scales peltate, blackish, with or without lighter, dentate margins; laminae 1-pinnnate 80-140 cm long; rachises stramineous, glabrous; basal pinnae 15-25 × 2.5-3.5 cm; costae abaxially and adaxially glabrous; veins commonly 1-furcate, rarely simple or 2 or 3-furcate (at the base of pinnae), abaxially and adaxially glabrous; sori marginal, discrete, born at the tip of single veins, placed side by side, opening extrorsely; outer indusia not modified; inner indusia hemi-umbonate, commonly forming wings connecting adjacent sori.

Geographical distribution: Endemic to the Brazilian Atlantic Forest, from Pernambuco to Santa Catarina.

Specimens examined: BRAZIL. Minas Gerais: Viçosa: Estrada São Miguel-Viçosa [Road to São Miguel], 25-IV-1930, Y. Mexia 4639 (F, IAN, MO-n.v., NY, PH, VIC, UC, US); Escola Superior de Agricultura e Veterinária, s.d., Y. Mexia 4964 (MO-n.v., NY, US, VIC); woods on slope, 710 m, 6-X-1930, Y. Mexia 5385-a (UC-2 sheets); Viçosa-Coimbra, km 120, Sítio Bom Sucesso, 13-IX-2002, G.E. Valente & M.L. Batista 1071 (VIC); Mata do Seu Nico, 20°47’S, 42°51’W, 800 m, 6-XI-2012, P.B. Schwartsburd & E. Guatimosin 2618 (NY, RB, UPCB, VIC).

Saccoloma elegans is recognized by the 1-pinnate laminae and the discrete sori placed side by side, opening extrorsely.

Acknowledgements

We thank Universidade Federal de Viçosa (UFV) and Programa de Pós-Graduação em Botânica; the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG) for the PhD scholarship to Nelson Túlio Lage Pena; the International Association for Plant Taxonomy (IAPT) for the “Grant 2020” to Nelson Túlio Lage Pena; the students and staff at Herbarium VIC; and Reinaldo Pinto for the illustrations.

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  • 1
    Part of the Doctoral Thesis of the first Author

Edited by

Associate Editor:

Regina Yoshie Hirai

Publication Dates

  • Publication in this collection
    28 July 2023
  • Date of issue
    2023

History

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