Sticherus bifidus (Willd.) Ching (1940: 282)

Lima, Lucas Vieira & Salino, Alexandre, 2018, The fern family Gleicheniaceae (Polypodiopsida) in Brazil, Phytotaxa 358 (3), pp. 199-234 : 212-214

3.1. Sticherus bifidus (Willd.) Ching (1940: 282) . Figs. 4A–E , 5A , 6A .

Mertensia bifida Willdenow (1804: 168) . Gleichenia bifida (Willd.) Sprengel (1827: 27) . Dicranopteris bifida (Willd.) Maxon (1909: 60) . Type:— VENEZUELA. Distrito Federal: Caracas, Bredemeyer s.n. (lectotype B [B-W-19468-01 0] photo!, designated by Proctor 1985, isolectotype S [S-R-3496] photo!).

Mertensia pubescens Humb. & Bonpl. in Willdenow (1810: 73). Gleichenia pubescens (Humb. & Bonpl. ex Willd.) Kunth (1815: 29) . Dicranopteris pubescens (Humb. & Bonpl. ex Willd.) Conzatti (1939:129) . Sticherus pubescens (Humb. & Bonpl. ex Willd.) Nakai (1950: 25) . Type:— VENEZUELA. Sucre: Cumaná, Santa Cruz, Humboldt & Bonpland s.n. (holotype B [B-W-19467-01 0]!).

Mertensia decurrens Raddi (1825:73) . Sticherus decurrens (Raddi) Gonzales in Gonzales & Kessler (2011: 23). Type:— BRAZIL. Rio de Janeiro: G. Raddi s.n. (lectotype PI, designated by Pic-Semolli 2005: 328, isolectotypes P [P00625735; P00625726] photos!).

Gleichenia mathewsii Hooker (1844: 9) . Mertensia mathewsii (Hook.) Fée (1866: 122) . Sticherus mathewsii (Hook.) Nakai (1950: 22) . Type:— PERU. Chachapoyas: Chachapoyas, 1844, Mathews 1092 pro parte (holotype K [K000589324]!).

Mertensia trifurcans Fée (1869: 201) . Gleichenia trifurcans (Fée) Christ in Schwacke (1900: 35). Sticherus trifurcans (Fée) Nakai (1950: 29) . Type:— BRAZIL. Rio de Janeiro: Tijuca, 8 April 1967, Glaziou 1697 (lectotype designated here, P [P00625775] photo!, isolectotype P [P00625774, P00625773] photos!, BR [BR0000006971748, BR0000006964191] photos!, S [S05-3159] photo!).

Plants terrestrial. Rhizomes 2.4–3.9 mm thick, with dark-brown, with scales narrow triangular, rigid, with apex acuminate, margins short to long ciliate. Fronds erect when young, becoming scrambling with age, 2–3(4)-forked,petiole 1.75–5.1 mm thick, ultimate branches 19–36.8 × 3.2–4.8 cm, lanceolate, apex pinnatifid, base truncate, rare slightly reduced, abaxial surface face dense to moderately scaly, rachis with scales light-brown, triangular, apexacuminate, base slightly cordate, margins short to long-ciliate, adaxial surface sparse to densely covered by hyaline filiform scales, segments linear 1.45–2.7 × 0.25–0.35 cm, margins plane to slightly revolute, abaxial surface dense to moderately covered by arachnoid scales on laminar tissue and secondary veins, midrib with arachnoid and smaller scales similar to the rachis. Buds with light-brown scales, with occasional apical or basal darkening, apex acuminate, base truncate, margins short to long-fimbriate, pseudoestipule present or not. Veins 1-forked. Sori medial, without paraphyses.

Distribution and habitat: — Brazil (Bahia, Ceará, Distrito Federal, Espírito Santo, Minas Gerais, Mato Grosso, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and São Paulo) Bolivia, Colombia, Costa Rica, Cuba, Ecuador, Guianas, Panama, Peru, Trinidad and Tobago, and Venezuela. In Brazil, this species occurs along roads and trails, and forest edge in the Atlantic forest domain at 150–1500 m.

Notes: — Sticherus bifidus is characterized by arachnoid scales present on the abaxial surfaces of the laminar tissue, secondary veins, segment midribs, and rachises; fimbriate bud scales; and mainly by the ciliate margins of the rhizome scales. Certainly, this is one of the most missapplied names in Gleicheniaceae , due to its large phenotypical plasticity and its close relationship to other species. Relative to the Brazilian species of the genus, S. bifidus is morphologically related to S. lanuginosus , S. paulistanus , and S. squamosus . It differs from S. lanuginosus by having branches pendent, larger segments (4 times longer than wide), fronds lesser forked, and by lacking of scales on the petioles. It differs from S. paulistanus and S. squamosus mainly by having bud scales fimbriate, arachnoid scales on the laminar tissue, rachises always terete, and rhizome scales ciliate.

Gonzales & Kessler (2011) recognized Sticherus decurrens as a different species from S. bifidus based in the apical bud scales darkening pattern, presence of pseudostipules, and fibrillose or arachnoid scales on abaxial surface of segments. In the present work, we chose to adopt a broad circumscription of S. bifidus because the differences listed by Gonzales (2003) and Gonzales & Kessler (2011) are quite variable between individuals from different populations, within the same population, or even in the same individual. Furthermore, Gonzales & Kessler (2011) recognized S. ferrugineus as a separate species from the S. bifidus complex, but, in that case, the Brazilian specimens identified as S. ferrugineus in several herbaria are in fact S. paulistanus (see notes on S. paulistanus ). These problems suggest that morphological studies do not satisfactorily resolve the circumscription problems in S. bifidus complex, and that future studies with different tools should be performed to test the monophylly of these proposed species and to provide precise species circumscriptions.

In the original description, Fée cited a collection of Glaziou (#1697) from Tijuca, Rio de Janeiro, Brazil. We found six exsiccatae of this Glaziou collection, three at P, two at BR, and one at S. Among the P specimens, we chose the one with original Fée label and in best conservation state. The BR specimens have transcript labels, but all information matches the protologe of the original description, as well the S material.

Selected specimens examined: — BRAZIL. Bahia: Arataca, Serra do Peito de Moça, estrada que liga Arataca à Uma , ramal ca. 22.4 km de Arataca com entrada no assentamento Santo Antônio. RPPN Caminho das Pedras, 15°10’25.0”S, 39°20’30.0”W, 1000 m, 15 February 2006, Matos et al. 968 (UPCB) ; Camacan, Fazenda Serra Bonita, 9.7 Km W de Camacã na estrada de Camacã para Jacarecí , daí 6 Km SW na estrada para a Reserva e Torre da Embratel , 15°23’30.0”S, 39°33’55.0”W, 835 m, 10 July 2005, Matos et al. 651 (UPCB). Ceará: Maranguape, Serra de Maranguape , Estrada Pirapora a Castelo, 27 June 1981, Martins s.n. (EAC10518). 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. 14890 (BHCB) ; Castelo, Parque Estadual Forno Grande , 1130 m, 25 June 2008, Salino et al. 13538 (BHCB) ; Divino de São Lourenço, Parque Nacional do Caparaó, RPPN Águas do Caparaó , 20º35’49.2”S, 41º46’52.2”W, 1000 m, 12 September 2008, Salino et al. 13847 (BHCB) ; Santa Tereza, Mata da Penha , 10 April 1984, Boone 42 (BHCB). Minas Gerais: Caeté , Serra da Piedade , 1600 m, 14 June 1997, Salino et al. 3142 (BHCB) ; Juiz de Fora, Campus Universidade Federal de Juiz de Fora, Faculdade de Direito , 21º46’22”S, 43º22’16”W, 885 m, 12 December 2014, Lima 90 (BHCB) ; Ouro Branco, Miguel Bournier , 20º25’49”S, 43º46’58”W 19 October 2009, Souza 978 (BHCB) ; Santa Rita do Itueto, Parque Estadual de Sete Salões, trilha da Gruta de Sete Salões e Pico do Garrafão a partir da fazenda dos Correa, 19º17’07”S, 41º22’29,6”W, 590 m, 09 May 2006, Salino et al. 11020 (BHCB) ; Simonésia RPPN Mata do Sossego , 20º04’02”S, 42º04’40,4”W, 1250 m, 20 May 2006, Salino et al. 11072 (BHCB). Paraná: Antonina, Reserva Natural Rio Cachoeira, Fazendo Rincão, 25º15’S, 48º41’W, 30 m, 1 March 2005, Matos 459 (UPCB) ; Guaratuva, estrada Alto da Serra- Ouro Fino, 29 June 1063, Hatschbach s.n. (PACA); Matinhos, Estrada Velha Alexandra , 11 August 1999, Dunaiki 1262 (UPCB) ; Paranaguá, Ilha das Cobras , 25°31’12.0”S, 48°30’33.1”W, 04 May 1986, Souza s.n. (UEC 67703) ; Tijucas do Sul, Vossoroca, 14 February 1974, Kummrow 344 (PACA). Rio de Janeiro: Cachoeiras de Macacu Street between Funchal and Guapiaçu, near REGUA, 22º28’73”S, 42°45’33”W, 37 m, 17 November 2009, Baber 447 (RB) ; Nova Friburgo, Distrito der Macaé de Cima , estrada para o Sítio Sophronitis , 25 October 1990, Sylvestre 339 (RB) ; Petrópolis, Mato do Judeo , 700 m, 07 December 1968, Sucre 4266 (RB) ; Rio de Janeiro, Parque Nacional da Tijuca, trilha para o Pico da Tijuca , 22º56’33.3”S, 43º17’11.1”W, 812 m, 7 March 2013, Mynssen 1387 (RB) ; Teresópolis Parque Nacional da Serra dos Órgãos, trilha Mozart Catão, 03 November 2004, Engelmann 042 (RB). Rio Grande do Sul: Osório, APA Morro da Borusia , 09, April 2008, Santos 68 (PACA) ; Pixirica , Morrinhos do Sul, 29°21’29.5”S, 49°56’48.0”W, 141 m, 10 September 2013, Gonzatti 866 (HUCS) ; Santo Antonio da Patrulha , 19 August 1993, Bueno et al. 3475 (PACA) ; São Francisco de Paula, Tainha, Serra do Pinto , 950 m, 5 May 2002, Lehn 345 (PACA) ; São Leopoldo, zona campestre, 20 May 2001, Oliveira, C.M. 121 (PACA). Santa Catarina: Blumenau, Parque Natural Municipal São Francisco de Assis , 26°55’17.4”S, 49°04’18.6”W, 25 April 1998, Hiendlmayer s.n. (FURB) ; Imaruí, Forquilha da Aratingaúba, Parque Estadual da Serra , 28°10’10.0”S 48°52’13.0”W, 604 m, 17 March 2010, Verdi et al. 4046 (FURB) ; Santo Amaro da Imperatriz, Cova da Onça , 27°37’44.0”S, 48°46’51.0”W, 236 m, 17 March 2010, Stival-Santos et al. 2111 (FURB) ; São Bento do Sul, Serra Alta , trilhos, rabo do macaco, 26°20’16.0”S, 49°23’01.0”W, 22 December 2013, Schwirkowski 149 (FURB) ; São Bonifácio, Rio Sete, 28°04’50.0”S, 48°57’39.0”W, 332 m, 12 April 2010, Verdi et al. 4370 (FURB). São Paulo: Eldorado, 6–10 km da Caverna do Diabo , 150 m, 09 September 1976, Davis 60658 (UEC) ; Itirapina, Próximo ao cerrado do Alvaro , 750 m, 06 July 1991, Salino 808 (UEC) ; Salesópolis, Estrada de Boraceia, represa do Rio Claro , s.d., Travassos 308 (RB) ; São Luiz do Paraitinga, Parque Estadual da Serra do Mar, trilha do Poço Pito , 23º18’42”S, 45º07’11.4”W, 900 m, 05 March 2001, Salino et al. 6203 (BHCB) ; São Miguel Arcanjo, Parque Estadual Carlos Botelho , 20 April 2002, Farias et al. 635 (UEC) .

Figures

FIGURE 4. A–E. Sticherus bifidus (Krieger s.n., CESJ 10201). A. Detail of the abaxial segment surface. B. Rhizome scale. C. Bud scale. D. Rachis scale. E.Arachnoid scale of the abaxial side of the laminae. F–I. S. brevitomentosus (Forzza et al. 7225, BHCB). F. Detail of the abaxial segment surface. G. Rhizome scale. H. Bud scale. I. Rachis scale. J–L. S. gracilis (Salino 3611, BHCB). J. Detail of the abaxial segment surface. K. Rhizome scale. L. Bud scale. M–R. S. holttumii (Almeida et al. 4174, BHCB). M. Detail of the abaxial segment surface. N. Detail of the abaxial segment surface. O. Bud scale. P–Q. Rhizome scales. R. Rachis scale.

FIGURE 5. Habit. A. Sticherus bifidus. B. S. brevitomentosus. C. S. gracilis. D. S. holttumii.

FIGURE 6. Distribution of five taxa of Sticherus in Brazil. A. Sticherus bifidus. B. S. brevitomentosus and S. holttumii. C. S. gracilis. D. S. lanuginosus.