Dicranopteris flexuosa (Schrad.) Underwood (1907: 254)

Lima, Lucas Vieira & Salino, Alexandre, 2018, The fern family Gleicheniaceae (Polypodiopsida) in Brazil, Phytotaxa 358 (3), pp. 199-234 : 201-204

1.1. Dicranopteris flexuosa (Schrad.) Underwood (1907: 254) . Figs. 1A–D , 2B , 3A , 12A–B and G .

Mertensia flexuosa Schrader (1824: 863) . Gleichenia flexuosa (Schrad.) Mettenius (1863: 50) . Type:— BRAZIL. Espírito Santo: Wied-Neuwied s.n. (lectotype designated here, BR [BR0000006870515], photo!, isolectotypes BR [BR0000006869731]; [BR0000006869823]; [BR0000006869793]; [BR0000006869762] photos!).

Gleichenia nitida Presl (1825: 70) . Mertensia nitida (C. Presl) Presl (1836: 51) . Dicranopteris nitida (C. Presl) Nakai (1950: 68) . Type:— MEXICO. Haenke s.n. (lectotype designated here, PRC [ PRC 45797] photo!, isolectotype PR [PR612503] photo!).

Mertensia rigida Kunze (1834: 16) . Dicranopteris rigida (Kunze) Nakai (1950: 69) . Type:— PERU. Chibangata: Poeppig 1153 (lectotype designated here, L [L-3537345] photo!).

Mertensia rigida Smith (1843: 381) , nomen nudum.

Mertensia scalpturata Fée (1869: 199) . Dicranopteris scalpturata (Fée) Nakai (1950: 69) . Type:— BRAZIL. Rio de Janeiro. Glaziou 1695 (lectotype designated here, P [P00633237] photo!, isolectotypes P [P00633238], [P00633239] photos!). Remaining syntypes:— BRAZIL. Claussen 102 (P [P00633240] photo!); Glaziou 364 (P [P00633236] photo!).

Plants terrestrial or epipetric. Rhizomes 1.8–4.3 mm thick, with red-brown rigid hairs, glabrescent. Fronds scrambling, 1–4-forked, ultimate branches 7.4–30 × 2–7 cm, linear to lanceolate, apex pinnatifid, base attenuate, ultimate segments linear 3.5–15 × 1.8–2.3 cm, margins slightly revolute, abaxial surface green or pruinose, glabrescent, with globose glandular hairs on the secondary veins. Buds covered by multicellular red hairs, pseudostipules present, accessory branches entire to pinnatisect. Veins 3–4-forked. Sori medial, without paraphyses.

Distribution and habitat: — Brazil (Alagoas, Amazonas, Amapá, Bahia, Ceará, Distrito Federal, Espírito Santo, Goiás, Maranhão, Minas Gerais, Mato Grosso do Sul, Mato Grosso, Pará, Paraíba, Pernambuco, Paraná, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina, Sergipe, São Paulo, and Tocantins), Bolivia, Central America, Colombia, Ecuador, Guiana, Mexico, Paraguay, Peru, Venezuela, and USA. It is a widespread species in Brazil that occurs between 100 m and 2000 m. It colonizes disturbed areas, where it may be dominant.

Notes: —This species has a broad distribution in Brazil, and the apparent absence of it in a few states (Acre and Piauí) may be due to lack of collection efforts in these areas. It occurs in all phytogeographic domains in Brazil, from sea level up to 2000 m Because of its broad distribution, this species presents great phenotypical plasticity, which results in a gradient of frond size, shape, and texture.

Epipetric specimens have significant blade reduction so that the fronds are deltoid and once-forked, usually with a long caudate apex. Because of the conspicuous differences in relation to terrestrial specimens, in herbarium determinations, the epipetric form of Dicranopteris flexuosa has repeatedly been misidentified for D. linearis . It does not have any taxonomic relevance, however, since this difference may be observed in the same individual with a portion terrestrial and other epipetric, the former with the traditional morphology, and the later with the reduced morphology described above.

Dicranopteris flexuosa is easily recognized by glabrous abaxial surface with only globose glandular hairs on the secondary veins, and presence of pseudostipules and accessory branches opposite to the main branches. It is morphologically related to Dicranopteris rufinervis , but this has whitish to red hairs on the midrib of ultimate segments on the abaxial surface, and coriaceous fronds. Glabrescent specimens of D. rufinervis may be mistaken for D. flexuosa , but have bacilliform glandular hairs on secondary veins rather than the having globose glandular hairs of D. flexuosa . In addition, D. flexuosa has the segment midribs strictly glabrous, whereas in D. rufinervis they bear multicellular hairs.

Several works that deal with the genus in Brazil assign Dicranopteris linearis (Burm. f.) Underw. (1907: 250) to the country (e.g. Barros & Xavier, 2009; Labiak & Prado 2003, Windisch, 1994). However, all the analyzed specimens previously identified as D. linearis are, in fact, D. flexuosa , D. rufinervis , or even D. seminuda . Furthermore, the distinctive characters used by those authors, like immerse veins and round ultimate segments apex, are not consistent due to the great phenotypical plasticity of the D. flexuosa . Mickel & Smith (2004) point out the resemblance between D. flexuosa and D. linearis , and they suggest that these names may be synonyms. In the preparation of the present work, a few specimens of D. linearis were examined and compared with Brazilian specimens of D. flexuosa . We concluded that it is impossible to distinguish unequivocally one from the other morphologically. Therefore, other studies must be performed to elucidate if there is a single species with Pantropical distribution or two separate species, one with Neotropical and the other with Paleotropical distribution. Phylogenetic studies using molecular data in the fern genus Rumohra Raddi (1819: 290) ( Bauret et al. 2017) demonstrate that, despite of morphologic uniformity of Rumohra adiantiformis ( Forster 1786: 82) Ching (1970: 34) , it actually represents several cryptic species and different phylogenetic lineages. Similar studies should be performed in Dicranopteris , aiming to investigate such possibilities and to elucidate part of its evolutionary history.

In the original description, Schrader described Mertensia flexuosa based on a Brazilian collection of Wied, but unfortunately, he did not mention the herbarium where the type collection was stored. Proctor (1985) cited that the type of M. flexuosa is in BR. However, in that herbarium there are five specimens of that species among Wied’s collection. Thus, we here proceed with a second step lectotype designation. Wied was in Brazil between 1815 and 1817 ( Moraes 2009), but the labels of BR exsiccatae date 1820 and 1829. However, there is a precedent of misunderstandings of Wied’s collections dates, and it seems that they actually represent the receiving date by Martius ( Moraes 2009, 2011). Therefore, such collections are likely to be the original material of M. flexuosa . Among the five exsiccatae, two have Schrader’s handwriting, one has Wied’s label, and the others have transcript labels. We chose the most complete specimen among the exsiccate with Schrader’s signature as lectotype.

Mertensia rigida was described by Kunze in 1834, as cited by the author, the type material was in the Poeppig- Kunze and Lehmann herbaria. Despite of a first locality indication by Kunze (“ Peru ad Chibangata Jul.1829 ”), other material from Brazil is cited as “see and determinate” in the Lehmann collection, but there is no holotype indication. This herbarium was partially incorporated by L. We here choose the specimen cited by Kunze in the protologue as lectotype since the type material storage in LZ was destroyed in WWII. Lellinger (1989) suggests that a possible isotype of G. rigida was in L, and here we confirm this information and designate it as the lectotype. Tryon & Stolze (1989) indicate type material in P, but this specimen was not found.

Gleichenia nitida was described based on a Haenke collection from Mexico, without number or herbaria specified. Haenke’s duplicates were distribuited to several herbaria, and we found two original specimens, one in PRC and another in PR. It is possible that other duplicates may be found in other herbaria. We chose as lectotype the PRC material, because it is the most complete specimen. Lellinger (1989), whithout seeing the type of G. nitida , considered this name as synonymous of Gleichenella pectinata , and were followed by Østergaard & Øllgaard (2001). However, we have examined the type material of G. nitida and it clearly corresponds to Dicranopteris flexuosa .

Mertensia scalpturata was described by Fée, and the original description lacks herbarium information. Besides that, three different collections are cited. We chose as lectotype the most complete specimen with Fée’s original label and signature, which is in P.

Selected specimens examined: — BRAZIL. Alagoas: Ibateguara, 29 November 1966, Pontual 305 (PEUFR); São José da Laje, Usina Serra Grande-Mata Maria Maior, 8°59’42.4”S, 36°07’28.9”W, 315 m, 08 February 2001, Pietrobom 4796 (UFP). Amapá: Porto Grande, 0º35’N, 51º44’W, 20 March 2001, Pereira 220 (BHCB). Amazonas: Manaus, estrada Manaus-Caracaraí, km 53, 27 September 1974, Conant 1124 (INPA); Manicoré, Parque Nacional do Campos Amazônicos, estrada do Estanho, 08º30’37”S, 61º36’10”W, 132 m, 01 August 2013, Almeida et al. 3310 (BHCB); Presidente Figueiredo, Barranco em beira da estrada de Balbina, próximo da chegada à Vila de Balbina, 01°00’S, 59°00’W, 25 September 2006, Zuquim 194 (INPA). Bahia: Coarci, Serra do Corcovado, próximo a propriedade do Sr. Vitorino, 14º42’12”S, 39º35’27”W, 550 m, 10 December 2014, Salino et al. 16026 (BHCB); Piatã, Serra da Tromba, 13°04’00.0”S, 41°54’00.0”W, 1339 m, 08 September 1996, Harley 28390 (HUEFS), Pilão Arcado, Brejo da Serra, Cabeceira da vereda de Brejo da Serra, 10°00’12.0”S, 42°30’15.0”W, 08 November 2009, Prata 2106 (ASE); Rio de Contas, Chapada Diamantina, Caminho para o Campo do Queiroz, 13°31’34.0”S, 41°54’18.0”W, 23 April 2011, Guedes 18863 (ALCB); São Sebastião do Passé, Recôncavo, Fazenda Panema, 12°32’48.0”S, 38°22’38.0”W, 25 March 2001, Loureiro 10 (ALCB). Ceará: Pacoti, perto de Guaramiranga, 4°13’30.0”S, 38°55’23.2”W, 720 m, 06 August 1944, Cutler 8153 (NY); Baturité, Subida da Serra de Baturite, 04 October 1990, Nunes s.n. (EAC). Distrito Federal: Brasília, Jardim Botânico de Brasília, Reserva Olho D’água. Coletado perto da Lagoa dos Sapos, 05 July 1999, Mascarenhas et al. s.n. (UB); Riacho Fundo, Fazenda Sucupira Região entre Recanto das Emas e Riacho Fundo, 15°51’52.0”S, 47°52’10.0”W, 1150 m, 17 September 1999, Rodrigues Júnior 1376 (HEPH). Espírito Santo: Cariacica, Reserva Biológica de Duas Bocas 20°18’13.9”S, 40°29’13.4”W, 425 m, 12 June 2010, Salino et al. 14917 (BHCB); Santa Teresa, Reserva Biológica Augusto Ruschi, trilha da Cachoeira, 19°55’14.1”S, 40°33’37.3”W, ca. 800 m, 02 December 2008, Salino et al. 14014 (BHCB); idem, 20°17’00.0”S, 40°28’12.0”W, 25 March 1990, Akahori s.n. (VIES); Viana, BR 262, 20°23’26.0”S, 40°28’58.0”W, 10 May 1988, Behar et al. 69 (VIES); Vila Velha, Interlagos, 20°19’47.0”S, 40°17’32.0”W, 21 December 2007, Souza et al. 78 (VIES). Goiás: Alto Paraíso de Goiás, Portal da Chapada, 14°09’52.8”S, 47°35’58.2”W, 1164 m, 06 October 2006, Rocha 63 (UB); Caiapônia, Serra do Caiapó, 48 km S of Caiapônia, 25 October 1964, Prance 59633 (UB); Catalão, Contraforte Central, ca. 26 km NE of Catalão Goiás, 875 m, 23 January 1970, Irwin et al. 25215 (UB); Ipameri, rod. GO-213, 23 January 1996, Pietrobom 2629 (MBM); Silvânia, 16°38’11.4”S, 48°38’50.5”W, 892 m, 06 December 2013, Elias 70 (UFG). Maranhão: Balsas, final da Chapada, 300 m, 24 November 1997, Oliveira et al. 737 (BHCB); Carolina, 7°07’48.0”S, 47°25’12.0”W, 14 April 1983, Taylor 1263 (NY). Mato Grosso do Sul: Corumbá, área da Mineração Urucum S.A., 29 June 1999, Assis et al. 27 (BHCB); Ponta Porã Fazenda Curupi, 02 March 2001, Carneiro 37 (BHCB); São Gabriel do Oeste, Reserva legal da Fazenda Bonito, 18º57’35’S, 54º18’17.4”, 280 m, 15 June 2002, Pott & Lima 5600 (BHCB). Mato Grosso: Barra do Garças, Distrito de Vale dos Sonhos, Serra de Vale dos Sonhos, 15°21’12.6”S, 52°13’32.9”W, 515 m, 22 February 2013, Athayde Filho et al. 4749 (NX); Campo Novo do Parecís, 13°02’00.0”S 57°57’00.0”W, 06 October 1996, Windisch 8469 (NX); Chapada dos Guimarães, Parque Nacional da Chapada dos Guimarães, Caminho para as cabeceiras do Rio Claro, 15º18’28”S, 55º51’28”W, 318 m, 01 March 2011, Almeida et al. 2669 (BHCB); Nova Xavantina, Estrada paralela ao bairro Deus e Amor, a cerca de 5 Km da cidade, ao redor de uma nascente chamada Olho d’água, 17 May 2009, Barbosa 9 (NX); Rondonópolis, Serra da Petrovina, A beira da rodovia, 400 m, 12 November 1989, Salino 634 (UEC). Minas Gerais: Açucena, margens do rio Corrente Grande, 18°56’58.6”S, 42°31’13.4”W, 470 m, 24 September 2009, Almeida 2105 (BHCB); Formoso, Parque Nacional do Grande Sertão Veredas, 15º11’30.8”S, 45º53’06.5”W, 760 m, 06 February 2006, Salino et al. 10758 (CESJ); Lima Duarte, Parque Estadual do Ibitipoca, Prainha, lado direito, subindo, 08 August 1993, Novelino & Yano 1170 (CESJ); Santa Rita do Jacutinga, Serra da Bandeira, 22°01’20.1”S, 43°59’39.8”W, 1620 m, 14 September 2014, Dittrich et al. 1916 (CESJ); Santana de Pirapama, Distrito de Fechados, 18°47’10.0”S, 43°52’26.0”W, 1100 m, 06 July 2009, Almeida 2025 (BHCB). Pará: Itaituba, Margens do rio Tapajós, a montante da sede do município de Itaituba, 4°22’54.0”S, 56°08’09.0”W, 25 m, 16 July 2016, Almeida 4338 (HSTM); Oriximinã, BR 163, estrada que dá acesso a perimetral Norte, a 14 Km de Cachoeira Porteira, 09 August 1986, Ferreira 7633 (INPA); Parauapebas, Serra Norte, 06°00’S, 50°18’W, 700 m, 24 May 1969, Cavalcante 2155 (MG); Rurópolis, Floresta Nacional do Tapajós, trilha que leva a Rio, 5 km após a comunidade de Piçarreiro (sentido Rurópolis), na altura do km 193 da BR-163 (Rodovia Cuiabá-Santarém), 3°56’17.0”S, 54°52’07.0”W, 116 m, 02 April 2016, Almeida 4280 (HSTM). Paraná: Adrianópolis, Parque Estadual das Lauráceas, 24°40’48.0”S, 49°00’00.0”W, 840 m, 11 January 2000, Dittrich 703 (NY); Curitiba, Parque Barigui, 26 March 1997, Kozera 546 (UPCB); Jaguariaíva, Parque Estadual do Cerrado, 24°10’S, 49°39’W, 800 m, 12 April 1994, Labiak 189 (UPCB); Ponta Grossa, rio Tibaji, 21 December 1971, Krieger s.n. (CESJ 11342); São José dos Pinhais, Usina Hidrelétrica de Guaricana, 11 July 1988, Straube 83 (MBM). Paraíba: Areia, Saburá, Margem da estrada para Alagoa Grande, 25 December 1986, Felix 478 (JPB); Mamanguape, Reserva Biológica Guaribas, Capim Azul. SEMA I, 6°44’30.7”S 35°08’31.4”W, 181 m, 15 February 1989, Santana 223 (JPB); Rio Tinto, Mata do Rio Vermelho, Fragmento Pb 113, 6°45’56.0”S 35°07’38.0”W, 94m, 29 March 2012, Gadelha Neto 3241 (JPB). Pernambuco: Água Preta, Engenho Sacramento, 8°41’08.2”S, 35°24’51.6”W, 229 m, 27 January 2000, Eugênio 453 (UFP). Rio de Janeiro: Itatiaia, 800 m, July 1933, Brade 12618 (RB); Nova Friburgo, Morro Alto do Teleférico, 23 July 1996, Pietrobom et al. 3372 (MBM); Petrópolis, subindo a Serra, próximo a rodoviária, 24 December 1992, Oliveira s.n. (CESJ); Rio de Janeiro, Parque Nacional da Tijuca, trilha para o Pico da Tijuca, 22º56’33”S, 43º17’11”W, 07 March 2013, Mynssen 1385 (RB); Santa Maria Madalena, Mata da Derrubadinha, 24 December 1977, Carauta 2763 (RB). Rio Grande do Norte: Nísia Floresta, Flona—IBAMA, 28 May 2003, Sanjuan 15 (MOSS). Rondônia: Colorado do Oeste, Linha 1, Km 32, rio Escondido, 20 January 2005, Neiva 81 (HBRA); Machadinho d’Oeste, Distrito de Tabajara, margens do Rio Machado, 08°55’27.0”S, 62°07’28.0”W, 84m, 02 June 2015, Labiak et al. 6133 (UPCB). Roraima: Amajari, estrada Boa Vista-Venezuela, Serra Pacaraima, 18 Km South of Santa Helena, 28 November 1977, Steward 218 (INPA). Rio Grande do Sul: Balneário Pinhal, horto florestal, 10 m, 13 October 2012, Gonzatti 618 (FURB); Pelotas, 08 June 1959, Sacco 1324 (UFRPE); São Francisco de Paula, 4 km de S. Francisco vindo de Taquara, 800 m, 10 March 1993, Windisch 8707 (NY); Torres, arredores do aeroporto, 15 April 2014, Gonzatti 1150 (FURB). Santa Catarina: Blumenau, Serra do Itajaí, Morro do Sapo, 27°03’39.7”S, 49°05’40.3”W, 312m, 18 December 2012, Funez 1144 (FURB); Indaial, Parque Estadual da Serra do Itajaí, estrada para sub-sede, 27°01’53.0”S, 49°09’55.2”W, 320 m, 22 April 2016, Funez 4685 (FURB); Rodeio, morro da Abissínia, 26º86’24”S, 49º36’22”W, 538 m, 18 March 2015, Funez 4219 (FURB). São Paulo: Bananal, Estação ecológica de Bananal, trilha até a sede, March 2001, Salino 6372 (BHCB); Brotas, Fazenda Rochedo, 06 May 1992, Lombardi s.n. (BHCB 20830); Caraguatatuba, Parque Estadual da Serra do Mar, estrada intermediária km 30, 23º41’32”S, 45º37’06”W, 600 m, 03 May 2001, Salino et al. 5299 (CESJ); São Paulo, March 1943, Krieger s.n. (CESJ 910B); Ubatuba, Parque Nacional da Serra do Mar, núcleo Pinciguaba, trilha do Picadão da Barra, 23º21’43.8”S, 44º50’02.3”W, 3 May 2001, Salino et al. 6710 (CESJ). Sergipe: Areia Branca, Serra de Itabaiana, Riacho Coqueiro, 10°44’52.0”S, 37°20’25.0”W, ca. 206 m, 11 July 2014, Santiago 1317 (ASE). Tocantins: Almas, região norte, Fazenda Minnehaha, arredores da antiga sede, ca. 70 km a nordeste da cidade de Almas, 11°08’18.0”S 47°07’20.0”W, 415 m, without date, Walter 5309 (CEN).

Figures

FIGURE 1. A–D. Dicranopteris flexuosa (Dittrich 2012, CESJ). A. Detail of the abaxial segment surface. B. Detail of the abaxial segment surface showing the globose glandular hairs on the secondary veins. C. Rhizome hairs. D. Bud hairs. E–H. Dicranopteris nervosa (Salino 6470, BHCB). E. Detail of the abaxial segment surface. F. Rhizome hairs. G. Bud hairs. H. Hairs of the abaxial segment surface. I–M. Dicranopteris rufinervis (Viveros 26, BHCB). I. Detail of the abaxial segment surface. J. Detail of the abaxial segment surface showing bacilliform glandular hairs on the secondary veins. K. Rhizome hairs. L. Bud hairs. M. Hairs of the abaxial midrib surface. N–Q. Dicranopteris seminuda (Rodriguez 3406, COL). N. Detail of the abaxial segment surface. O. Rhizome hairs. P. Bud hairs. Q. Hairs of the abaxial segment surface. R–U. Gleichenella pectinata (Gonzatti 1913, CESJ). R. Detail of the abaxial segment surface. S. Rhizome hairs. T. Bud hairs. U. Hairs of the abaxial segment surface.

FIGURE 2. Habit. A. Dicranopteris rufinervis. B. D. flexuosa. C. D. nervosa. D. D. seminuda. E. Gleichenella pectinata.

FIGURE 3. Distribution of five taxa of Gleicheniaceae in Brazil. A. Dicranopteris flexuosa. B. D. nervosa and D. seminuda. C. D. rufinervis. D. Gleichenella pectinata.

FIGURE 12. A–B. D. flexuosa (Almeida 302). A. Globose glandular hairs on the secondary veins of the segments. Scale bar 100 μm. B. Detail of a colapsed glandular hair, due to the herborization processes. Scale bar 25 μm. C–D. D. rufinervis. (Viveros 26) C. Baciliform glandular hairs on the secondary veins of the segments. Scale bar 150 μm. D. Detail of the glandular hair. Scale bar 50 μm. E. Spores of G. pectinata. F. Spores of S. salinoi. G. Spores of D. flexuosa. Scale bar 10 μm.