Ctenitis submarginalis (Langsd. & Fisch.) Ching (1940: 250) var. submarginalis

Viveros, Raquel Stauffer, Rouhan, Germinal & Salino, Alexandre, 2018, A taxonomic monograph of the fern genus Ctenitis (Dryopteridaceae) in South America, Phytotaxa 385 (1), pp. 448-450 : 448-450

23. 1. Ctenitis submarginalis (Langsd. & Fisch.) Ching (1940: 250) var. submarginalis . Figs. 10K , 26A–B , 27A .

Polypodium caripense Humboldt & Bonpland ex Willdenow (1810: 202) . Aspidium caripense (Humb. & Bonpl. ex Willd.) Mettenius (1856: 90) . Nephrodium caripense (Humb. & Bonpl. ex Willd.) Hooker in Hooker & Baker (1868: 265). Dryopteris submarginalis (Langsd. & Fisch.) Christensen f. caripensis (Humb. & Bonpl. ex Willd.) Christensen (1913a :

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Phytotaxa 335 (1) © 2018 Magnolia Press • 69 97). Ctenitis submarginalis f. caripense (Humb. & Bonpl. ex Willd.) Lellinger (1977: 710) . Type : — VENEZUELA. Cumaná, in umbrosis caripe, Humboldt 428 (holotype B–W 19700 -01 1! and B–W 19700 -01 2!, not isotype P 00669174!) .

Aspidium caripense (Humb. & Bonpl. ex Willd.) Mettenius f. macroloba Braun (1858 : 02). Type : — BRAZIL. Unknown s.n. (lectotype B 20 0068013!, designated here, isolectotypes B 20 0068010!, B 20 0068011!, B 20 0068012!).

Nephrodium tarapotense Hooker (1862: 107) . Dryopteris submarginalis (Langsd. & Fisch.) Christensen var. tarapotensis (Hook.) Christensen (1913a: 98) . Type: — PERU. Tarapoto, in Monte Campaña, Spruce 4016 (lectotype K 000200132! designated by Tryon & Stolze (1991), isolectotype P 01415576!); not isolectotypes: — in Monte Guayrapurina, Spruce 4016 (K 000200163!, P 00642705!).

Nephrodium crinitum Sodiro var. glaucescens Sodiro (1893: 251) . Dryopteris submarginalis (Langsd. & Fisch.) C. Chr. f. glaucescens (Sodiro) Christensen (1913a: 97) . Type: — ECUADOR. Sodiro s.n. (lectotype S-R-1777!, designated here, isolectotypes BM!, US 00067058!).

Nephrodium lagerheimii Sodiro (1893: 252) . Dryopteris lagerheimii (Sodiro) Christensen (1906: 273) . Dryopteris submarginalis (Langsd. & Fisch.) C. Chr. var. lagerheimii (Sodiro) Christensen (1913a: 98) . Type: — ECUADOR. Lagerheim s.n. (lectotype P 00642575!, designated here, isolectotypes C, P 00642674!, PH 00019282 [image!], SI 006581 [image!], SI 006582 [image!], S-R-1690!, UC 478166!, US 00067063!).

Dryopteris sellowii Hieronymus (1906: 324) . Type: — BRAZIL. Estancia Victoriano, March 1823, Sellow s.n. (lectotype B 20 0067944!, designated here, isolectotypes B 20 0067941!); the other syntype is: — BRAZIL. Rio Grande do Sul: São Leopoldo, Hamburguer, Stübel 1173 (B, not found).

Dryopteris collina Christ (1907: 922) . Type : — PARAGUAY. In sylvis collis Cerro, Hassler 513 (lectotype G 00349450!, designated here, isolectotype G 00307824 [image!]).

Dryopteris soriloba Christ (1909: 350) . Type: — PARAGUAY. Sierra de Amambay, in altaplanitie et declivibus, Rojas s.n. (Herb. Hassler 10454) (lectotype P 00642677!, designated here, isolectotypes BM 000511872!, G 00307823, G 00349449, G 00436365, MPU 015234); not Rojas s.n. (Herb. Hassler 10154—BM 000511873!, P 00642676!).

Selected specimens examined:— ARGENTINA. Corrientes : Santo Tomé, Ea. Timbol, 28 February 1983, Schinini et al.23565 ( UC) . Jujuy: Santa Bárbara, Abra de Los Morteros , 1400 m, 27 March 1992, Cabrera et al. 34853 ( NY) ; Misiones : Leandro N Alem, 10 March 1969, Krapovickas et al. 15096 ( UC) ; Libertador General San Martín, Predio UNLP, 27°05' S, 54°56' W, 19 July 1998, Biganzoli et al. 150 ( MO) ; Puerto Iguazú , 11 May 1951, Cabrera et al. 15 ( MO) ; San Antonio , 12 May 1951, Cabrera et al. 114 ( RB) ; San Ignacio, Santo Pipó , 05 April 1950, Schwarz 10367 ( RB) ; San Pedro, Caraguatay (centro), 11 May 1949, Montes 1668 ( RB) ; Salta: Rosario de la Frontera, 900 m, 28 July 1929, Venturi 9407 ( US); Tucumán: Famaillá, Caspinchango , 20 September 1949, Sas 144 ( NY) ; Santa Fé: Chaco Santafesino , December 1905, Venturi 239 ( GH) ; BOLIVIA. Chuquisaca: Tomina, Llantoj , 1400 m, 19°18'57" S, 64°05'15" W, 12 October 2004, Gutiérrez et al. 1017 ( MO) ; La Paz: Franz Tamayo, Sumpulo , 1180 m, 14°34'20" S, 68°46'38" 7 July 2008, Jimenez 5152 ( UC) ; Nord Yungas, Coroico, 1100 m, October 1912, Buchtien 3409 ( US); Luis Calvo: Serrania del Iñao , 1210 m, 19°29'47" S, 63°55'32" W, 4 October 2001, Huaylla 375 ( MO) ; Santa Cruz: Andrés Ibáñez , 840 m, 18°06' S, 63°30' W, 27 March 2002, Sundue & Nee 527 ( NY) ; Tarjita: Aniceto Arce Ruiz, Reserva Natural de Flora y Fauna Taríquia , 1540 m, 21°59' S, 64°32' 10 June 2004, Jimenez & 2404 ( NY) . BRAZIL. Alagoas: São José da Lage, Usina Serra Grande, Mata Maria Maior, Grota da Geraba , 380–415 m, 12 February 2001, Pietrobom & Santiago 5017 ( BHCB) ; Ceará: Baturité, Sitío Caridade , 6 December 1937, Eugenio 45 ( RB) ; Mulungu, Serra de Baturité , 800 m, 4°18' S, 38°59' W, 28 February 1997, Almeida Neto 104 ( HB) ; Goiás: Nova América , 16 March 1978, Gimontez s.n. ( RB) ; Mato Grosso: Caiapônia, Serra do Caiapó , 27 June 1966, Irwin et al. 17872 ( RB) ; La picada Guayra , August 1937, Cullen s.n. ( RB) ; Mato Grosso do Sul: Bonito, Gruta do Lago Azul , 500–600 m, 21º08’S, 56º28’W, 04 August 1994, da Silva & Rodrigues 1324 ( HB) ; Minas Gerais: Juiz de Fora, Rio Peixe , 13 March 1980, Krieger s.n. ( UEC) ; Monte Belo, Fazenda Lagoa , 19 January 1982, Boom 275 ( RB) ; Nova Lima, Estação Ecológica de Fechos , 20º03’57’’S, 43º57’32.7’’W, 11 July 2001, Salino et al. 7166 ( BHCB) ; Paraná: Capitão Leônidas Marques, Próximo ao Parque Nacional do Iguaçu , 15 August 2006, Labiak & Ribas 3732 ( RB) ; Cianorte, Forest Reserve of CMNP, 390 m, 2 April 1966, Lindeman & Haas 859 ( RB) ; Fênix, Parque Estadual de Vila Rica , 3 October 2008, Pereira & Falleiros 338 ( RB) ; Foz do Iguaçu, Parque Nacional do Iguaçu, Trilha da Represa , 185 m, 25º37’29.79’’S, 54º28’05.48’’W, 01 August

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2010, Viveros & Pastro 139 ( BHCB) ; Laranjeiras do Sul, Barra Rio Sossego , 4 January 1975, Hatschbach 35716 ( HB) ; Lobato, Fazenda Remanso Irmãos Ferraz , 22 July 1962, Gomes & Matos, 1035 ( RB) ; Rio Branco do Sul, Serra do Votuvoru , 20 April 1974, Hatschbach 34399 ( UPCB) ; Rio Negro , 18 December 1950, Frenzel s.n. ( RB) ; São Mateus do Sul , 20 February 1904, Gänsly 8 ( UC) ; Tomazina, Anhanguera , 2 September 1998, Hatschbach et al. 68286 ( RB) ; Toledo , 16 November 1963, Pereira 7911 ( HB) ; Turvo, Reserva Indígena de Guarapuava , 13 April 2002, Silva & Poliquesi 3619 ( RB) ; Rio de Janeiro: Duque de Caxias, Xerém , 22 March 1950, Brade & Duarte 20266 ( RB) ; Mangaratiba , Reserva Rio das Pedras, 6 January 2000, Mynssen 300 ( RB) ; Paraty , 11 May 1994, Marquete 1777 ( RB) ; Rio Bonito, Cachoeiras de Macacu , 25 m, 22°28'22" S, 42°45'39" W, 16 November 2009, Baber & Wesenberg KB 403 ( RB) ; Rio Grande do Sul: Canguçu , 20 June 1968, Cereni & Irgang s.n. ( ICN) ; Erechim, Esperança para Montenegro , 2 July 1949, Rambo 42340 ( RB) ; Esmeralda, Estação Ecológica Aracurí , 920 m, 18 September 1983, Cestaro s.n. ( ICN) ; Nova Petrópolis , 13 June 1949, Rambo 41961 ( RB) ; Passo Fundo , 20 February 1942, Mattos s.n. ( RB) ; Pelotas, 22 May 1959, Brauner 88 (F); Santa Tereza, Linha Pederneira - Reserva Brum , 7 January 2012, Gonzatti 297 ( RB) ; São Leopoldo, Morro das Pedras, 10 m, Dutra 479 (R); Santa Catarina : Araranguá , Morro do Meleiro , 18 November 1943, Reitz s.n. ( US) ; Caçador , Rio dos Bugres, 900 m, 23 April 1962, Reitz 12863 ( HB) ; Catanduvas , 28 December 1963, Reitz 16363 ( HB) ; Curitibanos, Ponte Alta do Sul , 900 m, 19 April 1962, Reitz 12583 ( HB) ; Itapiranga–Chapecó , 02 February 1951, Reitz 3792 ( RB) ; Mondaí, Linha Cascalho , 6 March 2009, Verdi et al. 1754 ( RB) ; Santo Amaro da Imperatriz, Vargem do Braço, Spannagel 448 ( US); São Miguel do Oeste , 1 March 1964, Castellanos 24769 ( HB) ; Xanxerê, 500–600 m, 25 December 1956, Smith & Klein 9201 (R); São Paulo: Bady Bassitt , Fazenda Nossa Senhora de Fátima , 500–600 m, 20°55' S, 49°27' W, 11 March 1995, Nonato 91 ( HB) ; Iguape , Serra dos Itatins, March 1924, Brade 8275 ( RB) ; Monte Alto, Serra Anhumas , 630 m, 20°22' S, 48°28' W, 3 June 1995, Pietrobom 1800 ( HB) ; Teodoro Sampaio, Parque Estadual Morro do Diabo , 400–600 m, 22°32' S, 52°11' W, 13 January 1995, Pietrobom 1519 ( HB) ; COLOMBIA. Antioquia: Amagá, 25 November 1937, Daniel 2221 (US); Magdalena: Santa Marta, Regnell 2581 (S); Risaralda: Pereira, Hacienda Alejandría , km 6 carretera La Virgínia-Cerritos, extremo norteño de parte ancha del Valle del Río Cauca , 11 February 1990, Silverstone-Sopkin 5893 ( UC) ; ECUADOR. Imbabura: 955 m, 0°10'19" S, 78°37'55", 24 August 2004, Croat & Ferry 93991 ( UC) ; Tungurahua: Baños Canton, Rio Verde , 1550 m, 01º24’S, 78º17’W, 07 July 1992, Fay & Fay 3557 ( MO) ; Zamora-Chinchipe: Yantzaza Cantón, along road out of the town of Chuchumbleza, pararell to Zamora River , going south, 800 m, 03º31’S, 78º31’W, 8 February 1993, Fay & Fay 4228 ( UC) ; PARAGUAY. Alto Paraná: Hernandarias, Reserva Biológica Itabó , 23 May 1989, Windisch 5468 ( HB) ; Puerto Presidente Strossner (currently Ciudad del Este ), km 12, Escuela Técnica Florestal, January-February 1982, Casas & Molero 5629 ( MO) ; Monumento Científico Moisés Bertoni , 25°39' S, 54°36' W, 21 July 1994, Zardini & Florentin 40022 ( UC) ; Amambay: Parque Nacional Cerro Corá , 18 September 1980, Foster s.n. ( UC) ; Caaguazú: Guayaquí, between Coronel Oviedo and Caaguazú, route 2, km 158–159, 25º29’S, 56º11’W, 26 August 1993, Zardini & Tilleria 37027 ( UC) ; Caazapá: Tavaí, Cerro Tupasy , 250–400 m, 29 October 1988, Soria 2421 ( MO) ; Canindeyú: Mbaracayú Natural Reserve , 24º 08’00’’S, 55º 31’41’’W, 13 January 1998, Zardini & Guerrero 47720 ( MO) ; Guairá: San Pedro, 15 December 1988, Soria 2931 ( MO) ; Itapúa: El Tirol, 19.5 km by road NNE Encarnación , 13 October 1981, Foster 81-13 ( UC) ; Paraguari: Macizo Acahay , 500 m, 25º54’S, 57º09’W, 16 June 1988, Zardini 4836 (MO, UC) ; PERU. Amazonas: Bongará, Sipabamba , 1300 m, 5 May 1981, Young & Eisenberg 322 ( UC) ; Cajamarca: San Ignacio, Distrito Huarango, San Martín- Quebrada Blanca , 900m, 5°19' S, 78°45' W, 15 May 1996, Campos et al. 2720 ( MO) ; San Martín: Huallaga, Saposoa , 1300–1350 m, 6°34'47" S, 77°23'06" 12 August 2000, Quipuscoa et al. 2058 (F); Rioja, Aguas Verdes, 900 m, 5°40' S, 77°40' W, 2 April 2001, van der Werff et al. 16614 (F); URUGUAY. Rivera: Rivera, 150–180 m, 1948, Herter s.n. ( MO); Tacuarembó: Erosionsschluchten im rotem Sandstein mit Urwaldresten, 20 March 1913, Ostern 6620 (S); Treinta y Tres: Isla Patrulla Quebrada de los Cuervos, March 1945, Rosengurtt 4826 ( GH); VENEZUELA. Distrito Federal: Los Venados, south side of Pico Avila, just north of Caracas, 12 September 1961, Tryon & Tryon 5726 (S); Tovar, 1854, Fendler 197 ( GH); Yaracuy: 850–990 m, 10°13'40" N, 68°37'00" 8 September 2001, Meier 8665 ( UC) .

Habitat and distribution:—Terrestrial in wet forests, 50–1600 m. United States of America (Florida and Lousiana), Mesoamerica ( Mexico, Guatemala, Honduras, Nicaragua, Costa Rica and Panama), West Indies (Hispaniola) and South America ( Venezuela, Colombia, Ecuador, Peru, Bolivia, Paraguay, Argentina and Brazil; Fig. 27A ; Tab. 01).

Notes:—The typical variety can be recognized by scales on costa abaxially lanceolate and with cordate base ( Figs. 26A–B ). Most specimens from Mesoamerica, West Indies and Andean countries are without indusia, or they

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Phytotaxa 335 (1) © 2018 Magnolia Press • 71 are small and inconspicuous. Nevertheless, most specimens from Paraguay, Argentina, Uruguay and southern Brazil have large and conspicuous indusia. The Andean specimens, from high elevations are usually larger and scalier plants, which scales can be darker than the most individuals from other Neotropical regions. The ploidy of individuals was not verified, and maybe differences of it can result such morphological variation.

Christensen (1913a) recognized the typical form of C. submarginalis as Dryopteris submarginalis f. vera , which does not constitute a validly published name (Art. 24.3 of ICN — McNeill et al. 2012). The other infraspecific taxa he considered were “f. tenuifolia ” (here treated as C. submarginalis var. tenuifolia ), “f. caripensis ”, “f. glaucescens ”, “var. tarapotensis ” and “var. lagerheimii ”, all these last ones are here considered as C. submarginalis var. submarginalis .

Willdenow (1810) did not cite any material when describing Polypodium caripense , just mentioned “habitat in America meridionali prope Caripe”. The original material consists of two sheets in B. Both correspond clearly to one single specimen, being one sheet the base of a lamina, at the top right is written “1a”, and the other sheet is the apical part of it, written “1b”. Both sheets are kept under one single cover, which label is written “Cryptogamia Filices, Polypodium caripense , frondibus bipinnatifidus integerrimis. Habitat Caripe”. Furthermore, only the sheet with the basal portion of lamina has the original label of Humboldt, a common practice when the specimen consists of more than one preparation kept together, but only the first is labeled. For this reason, those two sheets in B must be considered as the holotype (Art. 8.2, 8.3, Rec. 8A.4 of ICN — McNeill et al. 2012). Christensen (1913a) considered P. caripense as Dryopteris submarginalis f. caripensis , according to him this was an andine form, but occurring from Mexico to Venezuela. There is a sheet in P incorporated from Humboldt & Bonpland’s Herbarium, labeled as the collection 428. It consists of C. ampla and was probably erroneously labeled, then, must not be considered as an isotype.

Braun (1858) described Aspidium caripense f. macroloba based on a living plant cultivated in Horto Berolinense, originally from Brazil. There are four sheets of such original material in B not clearly labelled as being part of a single specimen (Art. 8.3 and 9.3 of ICN — McNeill et al. 2012). We designate as lectotype one sheet with the label handwritten by Braun. Christensen (1913a), considered this name under the “f. tenuifolia ”, which

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here we recognize as variety. However, all the original material of A. caripense f. macroloba , correspond, in our opinion, to C. submarginalis var. submarginalis .

Other one of the infraspecific taxon that Christensen (1913a) recognized in Dryopteris submarginalis was the other andine “f. glaucescens ”, based on Nephrodium crinitum var. glaucescens ( Sodiro 1893) . The examined original material of this name and other cited by Christensen (1913a) are densely covered with bacilliform trichomes between veins on both surfaces. The density of these or other kind of trichomes are quite variable in C. submarginalis . The abaxial surface is not glaucous as said by Sodiro (1893) and Christensen (1913a), or somewhat lighter than the adaxial. Among the original material (Art. 9.3 of ICN — McNeill et al. 2012), we selected for lectotype (Art. 9.12 of ICN — McNeill et al. 2012) a herbarium sheet in S, which is the most complete sample.

Sodiro (1893) also described Nephrodium lagerheimii , which Christensen (1913a) recognized as the andine form Dryopteris submarginalis f. lagerheimii . The original material and other from Andes at high elevations, are in general, the larger examined specimens, with larger, more numerous and somewhat darker scales. No holotype and no herbarium were specified in the protologue, some sheets of original material are in C, P, PH, S, SI and US (probably also in Q and QPLS). We selected a sheet in S as the lectotype (Art. 9.2, 9.5, 9.11, 9.12 of ICN — McNeill et al. 2012), the same was also seen, but not typified by Stolze ( Tryon & Stolze 1991).

Hieronymus (1906) described Dryopteris sellowii based on Stübel 1173 and Sellow s.n., both from southern Brazil. Although no herbarium was specified, it is probable that those collections were in Hieronymus’s herbarium, which was incorporated by B. Christensen (1913a) cited Stübel 1173 in B. However, no sheet with this information was found there, but two sheets of Sellow s.n. were, both with label handwritten “ Dryopteris sellowii n. sp. ” by Hieronymous. As lectotype (Art. 8.3, 9.2, 9.5, 9.11, 9.12 of ICN — McNeill et al. 2012), we chose the one with a small label written “Estancia Victoriano March 23”, as mentioned in the protologue.

Dryopteris collina was described by Christ (1907) based on the collection Hassler 513 from Paraguay. Hassler collections are mainly in G, where are found two sheets of Hassler 513 kept together, one is a leaf piece comprising the base of petiole to the middle of the lamina and the other is the middle to the apex. The first is with a photocopy of Hasssler’s original label (obviously included much later) and with another from “Herbier Delessert”. The second sheet is with Hasssler’s original label. So, the two sheets are otherwise not clearly labelled as being part of a single specimen, and should be interpreted as syntypes (Art. 8.3, 9.5 and 40 Note 1 of ICN — McNeill et al. 2012). Therefore, we designate here the sheet with the original label to be the lectotype.

Dryopteris soriloba , described by Christ (1909), was based on a collection in the Hassler’s herbarium, number 10454, collected in Paraguay by Rojas. Sheets of this collection are in BM, G, MPU and P. Christensen (1913a) cited this collection as type, in Christ’s and Roland Bonaparte’s herbaria. However, he starts his monograph informing that the material he examined from Christ’s herbarium were incorporated by Roland Bonaparte’s. This last, in turn, was incorporated by P. There are two sheets in P from Hassler’s herbarium written on the label “ Dryopteris soriloba Christ ”, “ type ” or “ spec. nov.”. One sheet has the stamp of Roland Bonaparte and is the 10454 of Hassler, while the other has the stamp of Christ’s herbarium, but its number is 10154, a probable original material not cited in the protologue. This can be a mistake of Christensen (1913a), once we did not find a sheet of Christ’s with the correct Hassler’s number. Thus, we attribute the lectotypification of Dryopteris soriloba to Christensen (1913a), although inadvertent (Art. 7.10, 9.9, 9.19 of ICN — McNeill et al. 2012).

Dryopteris sellowii , D. collina and D. soriloba were considered by Christensen (1913a) as D. submarginalis f. vera , the typical form of this species. The three names are based on specimens from southern South America ( Paraguay and the Brazilian state of Rio Grande do Sul) close to the type location of C. submarginalis var. submarginalis (the Brazilian state of Santa Catarina). The specimens from this area are somewhat morphologically constant.

23.2. Ctenitis submarginalis (Langsd. & Fisch.) Ching (1940: 250) var. tenuifolia (C. Presl) R.S. Viveros & Salino , comb. nov., stat. nov. Figs. 01C, 08A–B, 10L, 26C–D, 27B. Lastrea tenuifolia Presl (1849: 37) . Dryopteris submarginalis (Langsd. & Fisch.) C. Chr. f. tenuifolia (C. Presl) Christensen (1913a: 96) . Type:— BRAZIL. Rio de Janeiro, Pohl s.n. (lectotype PRC 454057! designated by Christensen 1913a, isolectotype BM!).

Phegopteris blanchetiana Fée (1952a: 245) , syn. nov. Type: — BRAZIL. Bahia. Blanchet 2928 (lectotype RB 00609504!, designated here).

Phegopteris oreopteridastrum Fée (1869: 97) . Type: — BRAZIL. Rio de Janeiro, Serra L´Estrella, Glaziou 963 (lectotype RB

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00543364!, designated here, isolectotypes C, P 00170062!, P 00170063!); remaining syntypes: — BRAZIL. Rio de Janeiro, Villa Rica, Vauthier 585 (P 00637536!, P 00637537!, P 00637538!, P 01415551!) .

Aspidium sancti-pauli Christ (1908: 21) . Type: — BRAZIL. São Paulo: Prope Fazenda Bela Vista in S. Cruz ad flumen Rio Pardo , Wettstein & Schiffner 2382 (lectotype P 00633040!, designated here).

Selected specimens examined:— BRAZIL. Alagoas: Ibateguara, Usina Serra Grande, Engenho Coimbra , Grota do Quirino , 390–450 m, 08º59’58.3’’S, 35º52’14.7’’W, 11 February 2001, Pietrobom & Santiago 4990 ( HB) ; São José da Lage, Usina Serra Grande , 380–507 m, 8°58'19" S, 36°06'36" W, 26 April 2002, Pietrobom 5483 ( HB) ; Ceará: Araripe, Base da Serra do Araripe , 08 August 1948, Duarte 1347 ( RB) ; Base da Serra do Arauju , 8 August 1948, Duarte 1347 ( RB) ; Baturité , Pereira s.n. ( US) ; Bahia: Camacan, Fazenda Serra Bonita , 835 m, 15º23’30’’S, 39º33’55’’W, 02 February 2005, Matos et al. 264 ( CEPEC) ; Feira de Santana, Fazenda Boa Vista - Serra de São José , 12°15' S, 38°58' W, 10 May 1984, Noblick 3169 ( MBM) ; Ilhéus, Centro de Pesquisas do Cacau , 28 October 1978, Mori 1100 ( CEPEC) ; Itabuna, Centro de Pesquisas do Cacau , 30 m, 14°45' S, 39°15' W, 10 January 1974, Harley 15011 ( CEPEC) ; Salvador, Parque de Pituaçú , 1 September 1997, Guedes 5202 ( MBM) ; Espírito Santo: Domingos Martins, Parque Estadual da Pedra Azul , na trilha para as piscinas naturais, 1470 m, 20º24’’08”S, 41º01’86”W, 5 December 2008, Salino et al. 14120 ( BHCB) ; Goiás: Aporé, Cachoeira do Corrente , 10 June 1993, Pietrobom 868 ( HB) ; Goiânia , December 1936, Brade 15365 ( RB) ; Serra do Caiapó, 900 m, 27 June 1966, Irwin et al. 17872 (F, NY); Jataí, Queixada, Margem do Rio Corrente , 09 February 1950, Macedo s.n. ( RB) ; Mato Grosso do Sul: Inocêncica , 400 m, 19º46’S, 51º48’W, 11 November 1995, Lucca Jr. et al. 116 ( HB) ; Minas Gerais: Caldas, Distrito de Pocinhos do Rio Verde , 1150 –1200 m, 21°56' S, 46°23' W, 16 June 1995, Pietrobom 1924 ( HB) ; Coronel Pacheco , 420 m, 9 August 1944, Heringer 1506 ( RB) ; Ouro Preto , Damazio, 442 ( OUPR) ; Viçosa, Agricultural College Lands , 680 m, 27 May 1930, Mexia 4753 ( RB) ; Passa Quatro , 3 May 1948, Brade 18940 ( RB) ; Paraná: Fênix, Parque Estadual de Vila Rica do Espírito Santo , 350 m, 23º54’54’’S, 51º57’13’’W, 09 November 2004, Labiak et al. 3451 ( RB) ; Morretes, 26 March 1914, Dusén 14732 (F); Rio Negro, Vila Nova , December 1904, Annies 75 ( RB) ; Pernambuco: Jaqueira, Usina Colônia , Mata do Ageró , 415 m, 08º44’27.4’’S, 35º50’37.7’’W, 18 October 2001, Lopes & Pietrobom 391 ( RB) ; Rio de Janeiro: Duque de Caxias, Xerém , 17 July 2003, Lemos s.n. ( RB) ; Itatiaia, Serra do Itatiaia , 18 June 1930, Brade 10065 (R); Mangaratiba, Reserva Ecológica Rio das Pedras, 19 October 1996, Braga et al. 3606 ( RB) ; Nova Friburgo, Morro do Porcelet , 1300 m, 19 July 1964, Pabst 8138 ( HB) ; Paraty, APA Cairuçu, Paraty-Mirim , 5–50 m, 23°01' S, 44°03' W, 7 March 1994, Marquete 1481 ( RB) ; Petrópolis , Spannagel 250 ( UC) ; Rio de Janeiro, Estrada da Vista Chinesa-Gávea , 8 May 1923, Hurgel 35 ( RB) ; Silva Jardim, Reserva Biológica de Poço das Antas, Trilha interpretativa do Projeto Mico Leão Dourado , 23 April 1997, Sylvestre et al. 1269 ( RB) ; São Paulo: Águas da Prata , Serra dos Poços, 850 m, 21°56' S, 46°48' W, 17 June 1995, Pietrobom 2001 ( HB) ; Campinas, Fazenda Sete Quedas , 20 November 1938, Viegas & Santoro s.n. ( RB) ; Cássia dos Coqueiros, Cachoeira Itambé, Rio Boiadeiro , 950–1000 m, 21°20' S, 47°08' W, 10 February 1997, Athayde 128 ( HB) ; Corumbataí, Serra do Padre , 700 m, 29 July 1993, Rodrigues 532 ( HB) ; Mendonça, Fazenda Santa Maria , 21°12' S, 49°35' W, 21 December 1996, Pietrobom 3868 ( HB) ; Monte Alto , September 1995, Lucca Júnior 52 ( HB) ; São Paulo, Morro do Jaraguá , 1000 m, 22 December 1912, Brade 5372 ( HB) .

Habitat and distribution:—Terrestrial. Endemic to Atlantic Forest, 200–1500 m. Northeastern to southern and central-west Brazil ( Fig. 27B ; Tab. 01).

Notes:— Ctenitis submarginalis var. tenuifolia differs from the typical variety by scales on costa abaxially filiform with truncate or rounded base ( Figs. 26C–D ). Like the typical, C. submarginalis var. tenuifolia is well represented in the herbaria and the presence of indusia is also an inconstant character. This taxon distribution is exclusively atlantic, sympatric with the typical variety, which is not only atlantic but widely distributed in the New World. Due to the sympatry, we recognized it under an infraspecific status.

The oldest name for this taxon is Lastrea tenuifolia described by Presl (1849). It was based on Pohl s.n. from Brazil. Mettenius (1858) and Christensen (1913b) confused it with Aspidium alsophilaceum ( C. aspidioides ), a name also based on a Pohl’s collection. Later, Christensen (1913a) realized and corrected this mistake. He also cited that the original material was in Presl’s herbarium. The collections of Presl’s herbarium are currently in PR or PRC (Presl worked in both). Only one sheet of Pohl s.n. with the name Lastrea tenuifolia is in PRC, and then we attribute the lectotypification to Christensen (1913a), although not intentional (Art. 7.10, 9.9, 9.19 of ICN — McNeill et al. 2012). A fragment of it is in BM (purchased from Christensen’s herbarium). There are three sheets of Pohl 3789 in W (W 0055893, W 0055894, W 0055895) kept as type of Lastrea tenuifolia and they seem

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to be parts of a single specimen from which fragments were also removed to compose the sheet in PRC. Then, their data may be more precise than what informed by Presl (1849) or, less probably, they are uncited original material.

Phegopteris blanchetiana was based on Blanchet 2928 from Bahia state in Brazil ( Fée 1852b). Christensen (1913a) included this name as an unknown species of Ctenitis . Such collection is in RB, but its label does not mention Bahia. In P there is a herbarium sheet (P00170082!), which on the label is written P. blanchetiana, Blanchet (without collecting number) and Bahia, so it seems to be at least an uncited original material. Both correspond to C. submarginalis var. tenuifolia . For lectotype, we designate the sheet in RB, which the collecting number matches with the protologue.

In the protologue of Phegopteris oreopteridastrum, Fée (1869) cited two collections: Glaziou 963 and Vauthier 585. Christensen (1913a) has already recognized this name under Dryopteris submarginalis f. tenuifolia and cited Glaziou 963 in Herbarium Hauniense (C), but without mentioning it as type. Then, we designate here a lectotype (Art. 9.12 of ICN — McNeill et al. 2012) for P. oreopteridastrum , the Glaziou 963 in RB, which contains the large and original Fée’s label with his signature, and the most complete sample.

When describing Aspidium sancti-pauli, Christ (1908) cited two localities. According to Art. 40 Note 2 of ICN ( McNeill et al. 2012) this does not constitute a specimen or gathering. The two localities are: Serra São João and Bella Vista. One original material is in P (from Roland Bonaparte’s herbarium) and on a label is written “ São Paulo. Prope Fazenda Bella Vista in district Urbis S. Cruz ad flumen Rio Pardo. Ca. 500 m. s. m. VII.1901, Wettstein & Schiffner ”, a different smaller label is with the number 2382, and has a Christensen’s identifying label from 1909. Other original material is in W (W 1905-0001039), on this there is a label written “ São Paulo. In sylvaticis “Serra São João” prope Santos. VIII.1901, Wettstein & Schiffner s.n. ”, but without a Christensen’s label. Certainly, they are the material to what Christ referred and both have a label with “ Aspidium Sancti Pauli n. sp. ” handwritten by Christ. Christensen (1913a) considered this name as synonym of his D. submarginalis f. tenuifolia and cited Wettstein & Schiffner 1901 in Christ’s herbarium and W. It seems that Christensen cited the collection year, not collecting number, but he was referring to those two specimens mentioned above. Then we designate here the sheet in P as the lectotype of A. sancti-pauli (Art. 9.12 of ICN — McNeill et al. 2012).

Names of Uncertain Application

Aspidium ameristonevron Fée (1857: 104) . Dryopteris ameristoneura (Fée) Christensen (1906: 251) .

Fée (1857) did not designate a type for this name and we did not find any specimen that we could ensure to be an uncited original material (Art. 9.3 of ICN — McNeill et al. 2012) in the herbaria where the American ferns which Fée described supposed to be. In the protologue, Fée (1857) presents only description and occurrence location ( Cuba). Christensen (1906) considered this taxon as Dryopteris , designating the new combination D. ameristoneura . Later, Christensen (1920) considered his combination as synonymous of D. grisebachii (here treated as C. grisebachii ). However, he cited D. ameristoneura not as a combination. Contradicting his previous work ( Christensen 1906), he demonstrated to be uncertain about the application of the Fée’s name, once he said that it was “probably not A. ameristonevron Fée ”. No uncited original material was found and the description of Fée is insufficient to diagnose which Cuban species he referred.

Aspidium caripense (Humb. & Bonpl. ex Willd.) Mett. f. brachyloba Braun (1858: 2) .

The description of Aspidium caripense f. brachyloba was based on a living plant from Colombia, cultivated in Horto Berolinense. We expected to find any herbarium sheet that could be considered as original material in B, were Braun worked. However, nothing that could be assumed as such was found in B, neither in other herbaria visited. We just can not ensure that an original material of it is extant, otherwise this name should be considered invalid (Art. 8.4 of ICN — McNeill et al. 2012). Christensen (1913a) has considered it as a synonym of Dryopteris submarginalis f. caripense . Probably it could correspond to C. submarginalis var. submarginalis , but as we have not examined any original material, and it seems that no one exists, we are not able to state that.

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Aspidium microcarpon Fée (1857: 105) , nom. illeg., non Blume (1828: 142). Type: — MEXICO. Córdoba, Schaffner 214 (not found).

Fée (1869) described three species of Ctenitis with laminae 1-pinnate-pinnatifid/pinnatisect based on Schaffner’s collections from Mexico: Aspidium microcarpon (Schaffner 214), A. obtusilobum (Schaffner 213) and A. microchlaena (Schaffner 459). Smith (1981), Tryon & Stolze (1991) and Mickel & Smith (2004) presumed that such collections were supposed to be in P. However, none of the three is in P. Fournier (1872), a French botanist who worked in P, cited them and he seems to be the last who have seen such materials (if he really saw). Ferns specimens described by Fée once belonged to Dom Pedro II, emperor of Brazil, but after the death of this monarch, they became the property of M. Cosson in Paris and later incorporated in P ( Underwood 1905). However, some Fée’s specimens remained in Brazil at Jardim Botânico do Rio de Janeiro, where is the RB herbarium. For example, the lectotypes designated here for Phegopteris blanchetiana (synonym of C. submarginalis var. tenuifolia ) and A. obtusilobum (synonym of C. microchlaena ). Schaffner 459 cited for A. microchlaena was found only in K, and we did not locate Schaffner 214 for A. microcarpon in none of the herbaria we have consulted (see Methods). Furthermore, the identity of this specimen is doubtful; according to the protologue, it could be either C. submarginalis var. submarginalis (with or without indusia) or C. microchlaena (always with indusia). Fournier (1872) considered A. microcarpon as the same of A. microchlaena , but distinguished them from A. obtusilobum . Christensen (1913a) without having seen the types, pointed that Dryopteris submarginalis f. caripensis (here recognized as C. submarginalis var. submarginalis ) corresponds to A. microchlaena and A. microcarpon as united by Fournier (1872) and treated C. microchlaena as D. karstenii . Probably guided by Fournier’s ideas, Mickel & Smith (2004) also considered A. microcarpon as synonym of C. microchlaena . There is a sheet in P (P01415596!) with a label written A. microcarpon , collected by Schaffner but without collecting number, from Orizaba, Mexico, differing from the protologue which informs “prés de Córdoba”. Maybe this could be an original material, however it is not with the usual Fée’s original label. Such specimen is C. submarginalis var. submarginalis , while the ones of A. microchlaena and A. obtusilobum correspond to C. microchlaena . Furthermore, the sori of this specimen are without indusia, which conflicts with the protologue. In the absence of a non doubtful original material, we are unable to ensure what Ctenitis species A . microcarpon represents.

Nephrodium vestitum var. squamigerum Mett. ex Baker (1870: 474) , nom. illeg. Type: — BRAZIL. Raddi s.n. (?), Martius 324 (P00170015!, B); Pohl s.n. (?); Milne s.n. (?); Bowie & Cunningham s.n. (?); Mac Gillivray s.n. (?); Glaziou 965 (P 00170004!, BR 0000013531638!, RB 00608023!), 2373 (P 00643972!, BR 0000006988357!, P 00643971!, P 01630479!), 2374 (P 00170005!, P 00642734!, P 01572335!); Lindberg 557 (B 20 0054405!).

Baker ( Hooker & Baker 1868) compared Nephrodium vestitum (here treated as synonym of C. deflexa ) to another species, Aspidium squamigerum , attributed to Mettenius [not ( Schlechtendal 1825: 23) Fée (1857: 104), not Mann (1868: 217)]. However, such name has never been published by Mettenius and it is not associated to a description or diagnosis by Baker ( Hooker & Baker 1868). Therefore, it is not effectively published (Art. 38.1 of ICN — McNeill et al. 2012).

In Flora Brasiliensis, Baker (1870) validly published Nephrodium vestitum var. squamigerum , specifying that such taxon referred to what Mettenius marked as A. squamigerum on specimens at Hooker’s Herbarium (currently in K). The types cited by Baker (1870) represent more than one taxon, e.g. Martius 324, Glaziou 965, 2373, 2374 correspond to C. deflexa , but Lindberg 557, corresponds to C. distans var. isabellina . Although more materials correspond to C. deflexa , the description is ambiguous and consequently it is not possible to be sure about which specimen (s) it refers the most nearly among the types cited. Besides that, Glaziou 965 was previously cited as type of Phegopteris fluminensis and 2373 of Aspidium basilare (Fée 1969) , both names that are lectotypified here (see C. deflexa header and notes). Thus, N. vestitum var. squamigerum is an illegitimate name (Art. 52 of ICN — McNeill et al. 2012).

Rosenstock (1904) treated Nephrodium squamigerum ( Rosenstock 1904: 224, not Hooker & Arnott 1841: 106) as a combination of a Mettenius’s name, which could be A. squamigerum . For this, Rosenstock included the vouchers Jürgens & Stier s.n. (J.-St. = Rosenstock exsicc. 32), Schmalz s.n. (S. = Rosenstock exsicc. 43), and Ulbricht s.n. (U. = Rosenstock exsicc. 55). We have not seen such specimens, but the short description matchs with C. deflexa .

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Christensen (1913a) included as synonym of Dryopteris ctenitis (here, C. distans ): A. squamigerum Mett. (referring to a note in Hooker & Baker 1868), Nephrodium squamigerum Rosenst. and N. vestitum var. squamigerum Baker (erroneously as Nephrodium caripense β squamigerum ). He recognized the infraspecific ranks D. ctenitis f. isabellina (here as C. distans var. isabellina ) and D. ctenitis f. amaurolepis (here as C. distans var. distans ). This last form he indicated that coincided with A. squamigerum of Mettenius. Like us, Christensen treated some of the types cited by Baker (1970). Lindberg 557 was cited by Christensen as D. ctenitis f. isabellina (to which we agree), and Martius 324 as D. ctenitis f. amaurolepis (to which we disagree).

Summing up, N. vestitum var squamigerum Mett. ex Baker was an attempt to interpret an unpublished name used by Mettenius and it is illegitimate due to its types were previously designated as types of other names. Even if it was an usable name, its types are composed of more than one taxon and the protologue does not allow knowing which specimen represents it better. Therefore, the application of this name remains uncertain.

Polypodium lachnopodium Smith (1846: 8) .

Smith (1846) made an enumeration of the ferns cultivated in the Royal Botanic Garden Kew. Among them, he described Polypodium lachnopodium as a new species based on a specimen from Jamaica introduced by Mr. William Purdie in 1843. No type was designated in the protologue, no original material was found in K, neither in other herbaria we have visited. It seems that the description was based only on a living plant, and as such, it is not acceptable as type (Art. 8.4 of ICN — McNeill et al. 2012). However, we can not ensure if an extant original material exists. Even son, Christensen (1920) suggested P. lachnopodium as a synonym of Dryopteris ampla ( Ctenitis ampla ). Nonetheless, the original description has few information to distinguish C. ampla from another decompound laminae species that occurs in Jamaica, e.g. C. grisebachii and C. excelsa (Desvaux 1827: 243) Proctor (1961: 34) .

Final Considerations

The Ctenitis distribution in South America here presented was expected according to literature data. It differs from other fern genera, which richness and endemism are in Andean regions. The Brazilian Atlantic coast has the highest species richness, mainly in the southeast and south as stated by Tryon & Tryon (1982b) and the northeast must be included in this range. Nonetheless, even though the Brazilian Amazonia (north) and the central-west are known for low richness of ferns, more collection efforts must be done, because those regions are still poorly represented and far from research centers.

Other perspectives for studies about Ctenitis could include cytogenetics. Species widely distributed as C. submarginalis var. submarginalis and C. ampla , are also morphologically variable, which could reflect variation in ploidy. Other feature that seems to influence the morphological variation is the elevation. At higher habitats, the individuals are scalier, and the scales are darker than the ones in lowlands. This was observed in C. ampla , C. falciculata , C. glandulosa , C. microchlaena and C. submarginalis var. submarginalis .

Most specimens of C. abyssi , C. aspidioides , C. fenestralis and C. laetevirens were with spores misshapen. This can suggest those taxa are hybrids, or their spores are malformed due to another reason. The fact is that they can be morphologically recognized and if hybrids, their parentage were not established.

The number of vascular bundles is an interesting feature in the major clades of ferns ( Smith et al. 2006), being constant in some families and genera. However, in Ctenitis this number is not constant. In South American Ctenitis it varies from 3–10. Why and how this evolved are remaining questions to be considered in evolutionary studies.

The recent Ctenitis phylogeny ( Hennequin et al. 2017) sampled 10 of ca. 50 New World species ( Viveros & Salino 2015) and so, much remains to be investigated, e.g. test the groups proposed by Christensen (1913a, 1920, 1934).

Aknowledgements

Our most sincere thanks to the curators and staff of the herbaria visited and those that lent a large amount of material. Special thanks to Alan R. Smith, Robbin C. Moran and Claudia Gonçalves for their assistance during the

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Phytotaxa 335 (1) © 2018 Magnolia Press • 77 visits; Belkiss R. Alméri for the line drawings; our colleagues, who accompanied us during fieldwork; Centro de Microscopia at the Universidade Federal de Minas Gerais; Coordenação de Aperfeiçoamento de Pessoal de Nível Superior ( CAPES) for the scholarships given to R.S. Viveros; Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the grants and financial support (especially to Reflora Program) given to A. Salino; Fundação Grupo Boticário de Proteção à Natureza for financial support for fieldwork; Marcelo Bueno for the richness map; Filipe S. de Souza for helping with distribution maps and images that allowed providing several illustrations; Sandra Knapp, Jefferson Prado, Jeferson M. Costa, Luís A.A. Góes-Neto and Elton L.M. de Assis for helpful comments and discussions about nomenclature; and finally, we thank João R. Stehmann, João A.N. Batista, Marcos E.G. Sobral, Vinícius A.O. Dittrich, Márcio R. Pietrobom, Adaíses S.M. da Silva and Claudine M. Mynssen for evaluating the doctoral thesis of the first author, from which this work is a result .

Figures

FIGURE 10. Spores of Ctenitis from South America. A: C. glandulosa (Salino 13626, BHCB, from Viveros & Salino 2015). B: C. grisebachii (Underwood 445, NY). C: C. laetevirens (Schimitt 561, BHCB). D: C. megalastriformis (Tuomisto 13393, UC). E: C. microchlaena (Mickel 2626, NY). F: C. nervata (Almeida 1682, BHCB). G: C. nigrovenia (Salino 15212, BHCB). H: C. paranaensis (Kozera 356, BHCB, from Viveros & Salino 2015). I: C. refulgens var. refulgens (Salino 15242, BHCB). J: C. refulgens var. peruviana (Schunke 5941, NY). K: C. submarginalis var. submarginalis (Verdi 1843, BHCB). L: C. submarginalis var. tenuifolia (Pietrobom 5015, BHCB). Scales bars = 10 µm.

FIGURE 26. Some morphological characters of Ctenitis from South America. A–B: Ctenitis submarginalis var. submarginalis (Verdi 1843, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: abaxial costa scale. C–D: C. submarginalis var. tenuifolia (Pietrobom 5015, BHCB). C: abaxial surface of a portion of medial pinna, showing indument. D: abaxial costa scale. Scale bars in A = 1 cm; B–D: = 1 mm.

FIGURE 27. Distribution of two varieties of C. submarginalis. A: C. submarginalis var. submarginalis. B: C. submarginalis var. tenuifolia.