Austroblechnum lehmannii (Hieron.) Gasper & V.A.O.Dittrich

Dittrich, Vinícius Antonio De Oliveira, Salino, Alexandre, Monteiro, Reinaldo & Gasper, André Luís De, 2017, The family Blechnaceae (Polypodiopsida) in Brazil: key to the genera and taxonomic treatment of Austroblechnum, Cranfillia, Lomaridium, Neoblechnum and Telmatoblechnum for southern and southeastern Brazil, Phytotaxa 303 (1), pp. 1-33 : 7-12

3. Austroblechnum lehmannii (Hieron.) Gasper & V.A.O.Dittrich in Gasper et al. (2016a: 203).

Blechnum lehmannii Hieronymus (1912: 473) . Blechnum lherminieri subsp. lehmannii (Hieronymus) Lellinger (2003: 146) . Type:— Colombia. Río Timbiqui, without date, F.C. Lehmann 8928 (holotype B 20 0031645, isotypes K000229679, K000229681, US 00067428, photo). Fig. 2A–C .

Blechnum mexiae Copeland (1932: 32) . Struthiopteris mexiae (Copeland) Ching (1940: 243) . Type:— BRAZIL. Minas Gerais: Carangola, 27 January 1930, Y. Mexia 4237 (holotype UC419577, photo, isotypes B 20 0031839, BM000769802, CAS284844, photo, K, NO0109674, n.v., NY, photo, P00627603, S, US 00067434, photo, VIC000016, photo).

Plants terrestrial; rhizomes ascending, long, non-stoloniferous, at apices densely clothed with atropurpureous to blackish, shiny, concolorous, narrowly triangular to lanceolate scales, 3.3–4 × 0.5–1.6 mm at the base, margins entire; fronds dimorphic, the fertile ones as long as the sterile, 23–33 cm long, the sterile 20–38 cm long; stipes completely stramineous or nigrescent proximally, with ovate, tan scales near apices and scales similar to those on rhizomes near bases, except for color (tan), of sterile fronds 3.5–8.7 cm long, 0.8–1.4 mm diam., of fertile fronds 5.5–13.1 cm long; sterile blades 18–28 × 4.1–5.7 cm, chartaceous, pinnatisect, narrowly elliptic, gradually reduced to apices and bases, proximal pinnae reduced to semicircular lobes or with lobes wider than long, glabrous; fertile blades 11–22 × 3.4–4.4 cm, pinnate, lanceolate to oblanceolate, truncate at base, with inconspicuous vestigial pinnae toward rhizomes; rachises glabrous on both sides or, mainly proximally, on the adaxial side, with tan, ovate-lanceolate scales; sterile pinnae 13–20 pairs, 2.2–3.2 × 1.2–1.8 cm at their bases, slightly to strongly ascending, fully adnate, triangular to broadly deltate, falcate, margins entire, plane or slightly revolute, apices acute, obtuse or rarely rounded; fertile pinnae 13–17 pairs, 19–35 × 1.8–2.7 mm, linear, strongly contracted; veins free, simple (distal ones), once forked (most) or 2 × forked (proximal pair), the proximal ones frequently arising from rachises, with hydathodes ending before margins.

Distribution and habitat:— Brazil (Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, and Rio Grande do Sul), furthermore Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, and Bolivia. This is a more or less common species in the study area, and therefore not threatened, growing generally in shaded areas close to streams in montane areas, in Tropical Atlantic Rain Forest, from (100) 300–1400 m.

Comments:— in the study area, the most similar species to A. lehmannii is A. squamipes . The first can be distinguished by the width of sterile fronds (> 4 cm, to 3.5 cm in A. squamipes ), the habitat (forest in A. lehmannii , grassland in A. squamipes ), the rhizomes ascending and long (> 20 cm in A. lehmannii —according to Sota (1972) to one meter long) or, in A. squamipes , erect to ascending, at most 5 cm long, the stipes of sterile fronds (stramineous or with a dark base in A. lehmannii , predominantly atropurpureous in A. squamipes ), the rhizome scales (narrowly triangular in A. lehmannii , ovate-lanceolate or ovate in A. squamipes ) and the absence (in A. lehmannii ) or presence (in A. squamipes ) of stolons. Young plants of Lomaridium plumieri (Desv.) C.Presl are also similar to A. lehmannii , but the scales of the former are linear and tan, generally with a dark central stripe, while in A. lehmannii these are narrowly triangular to lanceolate, atropurpureous to blackish and concolorous. Sehnem (1968) misapplied the name Blechnum onocleoides (Spreng.) Christ (sic) to members of this species in southern Brazil ( Blechnum onocleoides Swartz (1801: 75) ; according to Morton & Lellinger (1967), this is a synonym of Lomaridium fragile (Liebm.) Gasper & V.A.O.Dittrich , a species not found in Brazil. Mickel & Beitel (1988) proposed the synonymization of Blechnum lehmannii under Blechnum lherminieri (Bory in Kunze 1845: 173) Christensen (1905: 404) (= Austroblechnum lherminieri (Bory) Gasper & V.A.O.Dittrich in Gasper et al. (2016a: 203), stating that the plants are indeed the same species, with Lomaria lherminieri Bory being the older name. The senior author, however, has seen both type specimens at B and BM: A. lehmannii has narrower blades and, more importantly, the basal pinnae are gradually reduced to the rhizomes, while in A. lherminieri they are abruptly reduced. Considering these features, Lellinger (2003) reduced B. lehmannii to B. lherminieri subsp. lehmannii (Hieron.) Lellinger. We disagree with this decision and maintain A. lehmannii at species level. According to the photo presented by Kazmirczak (1999), the species exhibits vegetative propagation at the rhizome bases, a rare feature in the family and exclusive to this species in the study area.

Additional specimens examined:— BRAZIL. Minas Gerais: Alto Caparaó, Parque Nacional do Caparaó, Vale Verde, 1100 m, 09 October 1990, L. Krieger s.n. (CESJ 15071); Araponga, Parque Estadual da Serra do Brigadeiro, proximidades da sede, 1,400 m, 10 July 1999, A. Salino 4934 (BHCB, HRCB); Passa Quatro, Rio Retiro, 03 May 1948, A.C. Brade & E. Silva Araújo 18953 (K, RB); Simonésia, RPPN Mata do Sossego, 1,150 –1,600 m, 20º04’02’’S, 42º04’40’’W, 20 May 2006, A. Salino et al. 11070 (CESJ); Córrego da Lapa bei Ouro Preto, 23 May 1902, C.A.W. Schwacke 14597 (P). Espírito Santo: Castelo, Parque Estadual do Forno Grande, 20°31’51.1”S, 41°05’56.2”W, 1,450 m, 28 June 2008, A. Salino et al. 13707 (BHCB); Santa Teresa, trilha que sobe a encosta ao lado da entrada do Country Club, 25 February 1996, A. Salino 2638 (BHCB). Rio de Janeiro: Itatiaia, Serra do Itatiaia, Maromba, 25 June 1930, A.C. Brade 10200 (R); Macaé, Pico do Frade de Macaé, 900–1000 m, 22 October 1985, M. Leitman et al. 36 (RB); Nova Friburgo, Furnas, 17 March 1952, P. Capell s.n. (RB 77647); idem, 22°21’35”S, 42°34’00”W, 160 m, 01 September 2012, A. Bonnet 1600271 (FURB), Santa Maria Madalena, Parque Estadual do Desengano, 21°53’48”S, 41°54’04”W, 24 July 2012, C.M. Mynssen et al. 1314 (RB, photo); Teresópolis, Córrego Beija-flor, 1,200 m, 28 October 1929, A.C. Brade 9803 (R); idem, Serra dos Penitentes , 25 November 1975, R. Wels Windisch & A. Ghillány 409 (HB); 1874, A.F.M. Glaziou 9348 (K). São Paulo: Natividade da Serra, Parque Estadual da Serra do Mar, Núcleo Santa Virgínia, base de Vargem Grande, trilha para a Cachoeira da Boneca, 23°25’57”S, 45°12’36”W, ca. 800 m, 10 August 2001, V.A.O. Dittrich et al. 917 (HRCB); Pindamonhangaba, Fazenda São Sebastião do Ribeirão Grande, 28 November 1993, S.A. Nicolau & C.E. Espirito Santo 641 (SP); São Roque, 01 May 1977, R. Wels Windisch & A. Ghillány 651 (HB); Serra Negra , 03 June 1927, F.C. Hoehne s.n. (SP 20656); Campos da Bocaina, Invernada do Pinhal, 10 April 1894, A. Loefgren & G. Edwall s.n. (BM, SP 21674, SPF 94533). Paraná: Campina Grande do Sul, Serra Virgem Maria , 30 January 1969, G.G. Hatschbach 20959 (MBM, PACA); Morretes, Rio Sagrado de Cima, 100–300m, 08 August 1968, G.G. Hatschbach 19576 (MBM, PACA); idem, Jurapê, 08 February 1979, G.G. Hatschbach 41967 (MBM, PACA); idem, Parque Estadual Pico do Marumbi, ca. 600 m, 05 February 1999, V.A.O. Dittrich 589 (ICN). Santa Catarina: Araranguá, ca. 200 m, 07 December 1944, R. Reitz H655 (BM); Joinville, estrada Dona Francisca, 500 m, 21 June 1957, R. Reitz & R.M. Klein 4463 (B, HBR, MBM, PACA); idem, 26°11’33”S, 49°02’52”W, 576 m, 10 December 2009, T.J. Cadorin 933 (FURB); José Boiteux, Rio Laeisz, 26°54’43”S, 49°35’19”W, 823 m, 14 April 2010, A. Korte 2738 (FURB); Morro Grande, Três Barras, 28°42’36”S, 49°46’12”W, 311 m, 23 November 2009, M. Verdi 3146 (FURB); Paulo Lopes, Bom Retiro, 450 m, 13 December 1972, A. Bresolin 1053 (HBR, PACA); Rancho Queimado, Serra da Boa Vista, 1200 m, 04 February 1953, R. Reitz 5481 (HBR)—at PACA herbarium this number corresponds to Austroblechnum organense (Brade) Gasper & V.A.O.Dittrich ; São Bento do Sul, Braço esquerdo, 26°21’54”S, 49°14’08”W, 417 m, 21 November 2009, T.J. Cadorin 642 (FURB); Treviso, Nova Brasília, 28°26’37”S, 49°29’55”W, 555 m, 30 January 2010, M. Verdi 3533 (FURB). Rio Grande do Sul: Caraá, Fraga, Apa Municipal de Caraá, 30 November 2006, R.M. Senna & C. Mansan 1239 (HAS, photo); Maquiné, Reserva Biológica da Serra Geral , 05 February 2003, C. Bencke 531 (HAS, photo).

3. Austroblechnum organense (Brade) Gasper & V.A.O.Dittrich in Gasper et al. (2016a: 203). Blechnum organense Brade (1935: 2) . Lectotype (designated here):— BRAZIL. Rio de Janeiro: Serra dos Órgãos , 21 October 1929, A.C. Brade 9758 (R). Syntype: Brazil, Rio de Janeiro, Rio Roncador, 1750 m, 03 November 1929, A.C. Brade 9868 (R). Fig. 2D–F .

Plants terrestrial or epipetric; rhizomes erect or ascending, without stolons, at apices with lanceolate or narrowly triangular, dark tan to blackish, concolorous or not, translucent (the tan ones) or not (the other ones), shiny, 4.7–6 × 1.5–1.8 mm at the base, margins entire; fronds dimorphic, the fertile ones longer, shorter or as long as the sterile fronds, 31–50 cm long, the sterile 27–36 cm long; stipes atropurpureous or stramineous, sometimes atropurpureous and stramineous proximally (on different sides), with scales only near the rhizomes, these like those on rhizomes, lacking hairs, of sterile fronds 7.2–11.3 × 0.9–1.9 mm; of fertile fronds 21 cm long; sterile blades 19–23 × 6–11.6 cm, ovate or lanceolate, pinnatisect, the apices gradually reduced, truncate at base, with 0–1 reduced pinnae pairs and (0)1–3 pairs of vestigial pinnae, glabrous; fertile blades 9.3–13.6 × 6.4–10.8 cm, pinnate, ovate or deltate, truncate at base, with 2–4 pairs of almost imperceptible vestigial pinnae; rachises glabrous on both sides; sterile pinnae 9–19 pairs (excluding the vestigial ones), 3.2–8.4 × 0.8–1.7 cm, generally ascending, fully adnate, slightly falcate, apices acute to acuminate; veins free, once forked at proximal and median parts, distally simple, arising from costae or from rachises, with hydathodes at the margins or before.

Distribution and habitat:— Brazil (Minas Gerais, São Paulo, Santa Catarina, and Rio Grande do Sul [first record]), furthermore Colombia and Venezuela [first record]). This species occur in areas between 500 and 1700 m a.s.l., on plateaus and in mountain ranges such as Serra do Mar and Serra da Mantiqueira , generally close to streams in deep shade of forests. In southern Brazil it can be found from 500 to 1200 m, whereas in southeastern Brazil it is confined to higher elevations (above 1300 m).

Comments: —this species is very close and maybe even conspecific with A. divergens . The features distinguishing both can be seen under A. divergens . The species is also similar to Cranfillia mucronata (Fée) V.A.O.Dittrich & Gasper , from which it can be distinguished by the size and form of rhizome scales and by the presence of vestigial pinnae at the base of blade in A. organense (no vestigial pinnae in C. mucronata ). Brade (1935) did not designate a holotype in his description of Blechnum organense , citing two syntypes (A.C. Brade 9758 and 9868). Both specimens are in good condition, but 9758 is in better state and is therefore designated as lectotype.

Additional specimens examined:— BRAZIL. Minas Gerais: Delfim Moreira, 22°36’35”S, 45°20’30”W, 175 m, 17 March 2011, A.L. de Gasper 2658 (FURB); Marmelópolis, picada para o pico dos Marins, entre 22°30’ e 22°31’S e 45°08’30” e 45°09’30”W, 1,450 m, 03 April 2002, V.A.O. Dittrich 1122 (HRCB); Rio de Janeiro: Teresópolis, Serra dos Órgãos , córrego Roncador, 1,700 m, 15 July 1940, A.C. Brade 16373 (K, RB); ibidem, córrego Beija-flor, 1,300 m, 16 August 1940, A.C. Brade 16533 (K). São Paulo: Campos do Jordão, Parque Estadual de Campos do Jordão, trilha da Cachoeira da Celestina, ca. 22°42’S, 45°28’W, 1,350 –1,500 m, 27 November 2001, V.A.O. Dittrich & A. Mantovani 1067 (HRCB). São José do Barreiro, Bocaina, Fazenda do Lageado, 1,600 m, March 1951, F. Segadas Vianna & M. Starling 2661 (R). Unknown municipality: Serra da Bocaina , 1,700 m, 09 May 1957, A.C. Brade 20911 (K, RB). Santa Catarina: Rancho Queimado, Serra da Boa Vista , 1,200 m, 04 February 1953, R. Reitz 5481 (PACA) (at HBR herbarium this number corresponds to Blechnum lehmannii Hieron. ). Rio Grande do Sul: Caraá, Fraga, APA municipal de Caraá, 03 December 2008, R.M. Senna 1343 (HAS). VENEZUELA. Colonia Tovar, 1856, J.W.K. Moritz 455 (B).

4. Austroblechnum penna-marina (Poir.) Gasper & V.A.O.Dittrich in Gasper et al. (2016a: 203). Blechnum penna-marina (Poir.) Kuhn (1868: 92) . Polypodium penna-marina Poir. in Lamarck (1804: 520). Lomaria penna-marina (Poir.) Trevisan (1869: 570) . Struthiopteris penna-marina (Poir.) Maxon & Morton (1939: 43) . Type:— CHILE, Straits of Magellan, P. Commerson s.n.. (holotype P, photo, isotypes FI, BM000769833, photo). Fig. 3A–C .

Blechnum penna-marina var. boliviana Rosenstock (1913: 60) . Blechnum penna-marina (Poir.) Kuhn subsp. boliviana (Rosenst.) Chambers & Farrant (1996: 98) . Type:— BOLIVIA, Unduavi, Nordyungas, 3,300 m, November 1910, O. Buchtien s.n. (Fil. Bol. Exsic. 45) (holotype BM000769834, isotype P).

Lomaria polypodioides Gaudichaud ex Desv. (1825: 98) (non Blechnum polypodioides Raddi (1819: 294) , nec Blechnum polypodioides ( Swartz 1788: 127) Kuhn (1868: 92) = Lomaridium fragile ( Liebmann 1849: 232) Gasper & V.A.O.Dittrich in Gasper et al. (2016a: 212). Type:— Falkland Islands, East Falkland ( Isla Soledad), without date, J.S.C.D. d’Urville s.n. (holotype P n.v., isotype FI).

Plants terrestrial; rhizomes long-creeping, stoloniferous, the scales dark tan, concolorous, narrowly triangular, lanceolate or ovate, 1.5–2.5 × 0.7–0.8 mm at the base, margins entire; fronds dimorphic, the fertile ones longer and narrower than the sterile, 16–51 cm long, the sterile 8.1–37 cm long; stipes atropurpureous or light red, clothed mainly proximally with tan to light tan, concolorous, triangular or ovate scales with entire margins, longer on fertile fronds, of sterile fronds 0.5–11.6 cm long, 0.9–1.5 mm diam., of fertile fronds 19–27 cm long, wholly atropurpureous or stramineous distally; sterile blades 7.3–21 × 1–3.3 cm, subcoriaceous, pinnatisect or sometimes proximally pinnate, with sparse multicellular hairs on abaxial side, mainly on costae but also on veins, oblanceolate, pectinate, gradually reduced to apices and base, proximally the pinnae wider than long; fertile blades 14–30 × 1.2–3.3 cm, pinnate, narrowly obtrulate, gradually or more or less abruptly reduced proximally with some vestigial pinnae toward the rhizomes; rachises clothed, mainly proximally, with lanceolate, light tan scales abaxially and, adaxially with sparse, linear (with expanded bases), tan, concolorous scales; sterile pinnae 19–48 pairs, 0.5–1.7 × 0.2–0.6 cm, slightly ascending to slightly descending, more frequently patent, fully adnate, narrowly oblong, margins entire, slightly to strongly revolute, apices obtuse or rounded; veins free, once forked, rarely some distal veins simple, terminating as hydathodes at the margins.

Distribution and habitat:— Brazil (Minas Gerais [first unequivocal record], Rio de Janeiro, São Paulo, Santa Catarina, and Rio Grande do Sul), furthermore Peru, Bolivia, Argentina, Chile, Australia, New Zealand, and southern islands of Atlantic (e.g. Falkland Islands, Tristan da Cunha), Indian and Pacific oceans. This species is not very common in Brazil, being restricted to elevated, cold, open areas mainly in the south of country (southernmost plateaus, close to Serra Geral escarpments), but also on Itatiaia plateau (Mantiqueira range), above 800 m in south and from 1650–2350 m at the Itatiaia massif (southeast). It is not considered threatened ( Dittrich et al. 2013).

Comments: —Looser (1947) apud Kazmirczak (1999) accepted four varieties in this species, based mainly on quantitative characters and blade texture. Chambers & Farrant (1996) also split the species into four taxa, but at a different rank (subspecies). These authors also used quantitative characters in circumscribing these subspecies, although they considered stipe color and shape of pinna bases and pinna margins important features. Sota (1970) studied the taxonomy, ecology, and distribution of this species thoroughly, but did not subscribe to importance of the characters used by Looser (and later by Chambers & Farrant); consequently, he chose not recognize infraspecific taxa, since the morphological variation he observed was clinal and ecologically dependent. We agree with Sota’s (1970) observations (1970) and, accordingly, we prefer not to recognize infraspecific taxa.

Additional specimens examined:— BRAZIL. Minas Gerais: Itamonte, Parque Nacional do Itatiaia , a caminho de Vargem Grande, 22°20’16,9”S, 44°44’6,6”W, 12 July 2007, A. Salino et al. 12493 ( BHCB). Rio de Janeiro: Itatiaia, Serra do Itatiaia, ca. 2,350 m, June 1902, P.K. Dusén s.n. (K) ; idem, Serra do Itatiaia , 20 June 1930, A.C. Brade 10090 ( RB) ; Resende: a ca. 1 km do Abrigo Rebouças , ca. 2,200 m, 24 July 2004, V.A.O. Dittrich & M.C.O. Jorge 1313 ( HRCB). July 1874, A.F.M. Glaziou 7324 (K) ; August 1877, A.F.M. Glaziou 9059 (K). São Paulo: Campos do Jordão , 20 September 1921, F.C. Hoehne s.n. ( SP 8699 , SPF 94523 ) ; idem, April 1937, L. Lanstyack s.n. (K, RB 33131 ). Santa Catarina: Bom Jardim da Serra, Morro da igreja , 28°07’24”S, 49°28’48”W, 08April 2010, A. Salino 14745 ( FURB) ; Bom Retiro , Campo dos Padres, 16 December 1948, R. Reitz 2379 ( RB) ; Lages , 1905, C. Spannagel s.n. (Fil. Austr. Exsic. 236) (K) ; idem, 1921, C. Spannagel 84 ( HB, SP) ; São Joaquim, Morro da Igreja , 1,822 m, 22 January 1960, J. Mattos 7032 ( PACA) ; idem, near encruzilhada de Boava, 8 km south of São Joaquim , ca. 28°20’S, 49°56’W, 1,300 m, 05 January 1965, L.B. Smith & R. Reitz 14286 ( HBR, R) ; idem, rodovia de acesso a São joaquim, capão de floresta, 28°21’39”S, 49°58’16”W, 09 April 2010, A. Salino et al. 14786 ( FURB) ; Urubici, Parque Nacional de São Joaquim , 28°08’43”S, 49°37’06”W, 1,384 m, 09 May 2009, A.L. de Gasper 2090 ( FURB). Rio Grande do Sul: Bom Jesus, without date, J. Dutra 746 ( PACA) ; idem, Serra da Rocinha , 1,000 m, 19 January 1950, A. Sehnem 4324 ( PACA) ; Cambará do Sul, Itaimbezinho , 19 December 1950, A. Sehnem 5203 ( MBM, PACA) ; São Leopoldo, 10 October 1934, A. Sehnem s.n. ( SP 50620 ). Unknown municipality: Rio dos Touros, Aparados da Serra , 800 m, 13 January 1942, A. Sehnem 948 ( PACA). Unknown state: Resende (Rio de Janeiro) or Itamonte (Minas Gerais), 16 February 1981, M. Ranal 175 ( HUFU) .

Figures

FIGURE 2.A–C. Austroblechnum lehmannii.A. Habit. B. Pinna and venation pattern.C.Rhizome scale.D–F.Austroblechnum organense. D. Habit. E. Pinna pair showing venation. F. Rhizome scale. (A–C from A. Salino et al. 14054, BHCB; D–F from A.L. Gasper et al. 2658, BHCB).

FIGURE 3. A–C. Austroblechnum penna-marina. A. Habit. B. Pinnae and venation pattern. C. Rhizome scale. D–F. Austroblechnum squamipes. D. Habit. E. Base of blade showing pinnae and venation. F. Rhizome scale. (A–C from A. Salino et al. 14745, BHCB; D–F from A. Salino et al. 12036, CESJ).