Aspidium ctenitis Link (1833: 122) Nephrodium ctenitis (Link) Baker Hooker & Baker (1868: 265) Dryopteris ctenitis (Link) Kuntze (1891: 812) Aspidium amaurolepis Fée (1869: 137) Dryopteris ctenitis (Link) Kuntze f. amaurolepis (Fée) Christensen (1913a: 94) A taxonomic monograph of the fern genus Ctenitis (Dryopteridaceae) in South America Viveros, Raquel Stauffer Rouhan, Germinal Salino, Alexandre Phytotaxa 2018 2018-01-23 385 1 448 450 6BN8P (Brack.) Ching (1938: 277) Ching 277 1938 Brackenridge 1854 [179,667,547,573] Polypodiopsida Dryopteridaceae Ctenitis Plantae Polypodiales 37 Tracheophyta species distans   Stemserect or ascending, 1.5–3.5 cmdiam., scales 4.9–16.1 × 0.2–0.9 mm, light castaneous, clathrate or subclathrate, linear-lanceolate or lanceolate, entire, with or without some short fimbriae at base; leaves(34) 78–187 cmlong; petioles(14) 28–71 cm× (1.7) 3.0–6.0 mm, with 4 or 6 vascular bundles at base, stramineous, scales 3.0–7.4 × 0.2–0.6 mm, light castaneous on petiole base, becoming dark brown to blackish on distal portion, clathrate, not tangled on petiole base, becoming appressed towards distal portion, flattish, flaccid, linear-lanceolate or lanceolate with cordate base and filiform apex, denticulate, with or without some short fimbriae at base and laterally, sparse or dense catenate trichomes abaxially, glandular trichomes absent; laminae(20) 50–116 × (14) 22–46 cm, width ca. 1/3–1/2 of length or wider, 1-pinnate-pinnatisect or 1-pinnate-pinnatifid basally, 1-pinnate-pinnatifid medially and apically, lanceolate or ovate, apex confluent; rachisesstramineous, scales like those on distal portion of petioles, rare to dense catenate trichomes abaxially, glandular trichomes absent; pinnae10–37 pairs, the basal and medial ones stalked to 6.7 mmlong, the apical ones sessile, basal pinnae basiscopically and acroscopically somewhat equally developed, the medial 7.5–22.5 × 1.8–4.6 cm, lanceolate, incised more than 3/4 of the distance between the segment apex and costa, basal segments somewhat shorter or as long the next, apex attenuate; adaxial pinnae axeswith sparse scales on costa, 0.6–3.1 × 0.06–0.12 mm, dark brown to blackish, filiform, catenate trichomes dense on costa, sparse on costule, rare or sparse on veins, bacilliform trichomes absent; adaxial laminar surface between veinsglabrous; abaxial pinnae axeswith sparse to dense scales on costa and costule (basally), (0.6) 1.2–4.3 × 0.1–0.7 mm, mainly castaneous, but dark brown to blackish towards apex or uniformly dark brown or blackish, clathrate or subclathrate, ascending, sometimes imbricate, mostly flattish, but can be vaulted at base, flaccid, lanceolate or linear-lanceolate with cordate or rounded base and filiform apex, denticulate (the teeth claw-shaped), with or without some short fimbriae at base and laterally, proscales 0.6–3.1 mmlong sparse or absent on costule, catenate trichomes absent or sparse on costa, costule and veins, bacilliform trichomes absent or sparse on costule and veins, glandular trichomes absent, filiform trichomes absent; abaxial laminar surface between veinsglabrous or with sparse bacilliform trichomes, or sparse catenate trichomes; segments19–27 pairs, 2.8–5.6 mmwide, patent or subfalcate, entire, repand or serrate towards apex, apex rounded or acute, margin with catenate trichomes, the distance from each other is the same width of the segments or narrower; veinssimple, 7–15 pairs per segment, the basal ones from adjacent segments end at margin well above the sinus; soriinframedial, indusia conspicuous, entire or ciliate, with bacilliform trichomes; sporeswith coarse folds and large tubercles.  Notes:—The specimens of  C. distansare usually large and can be recognized by the dark brown to blackish scales on rachis and, castaneous but dark brown to blackish towards apex on costa and costule abaxially, or uniformly dark brown to blackish. These scales are denticulate, which teeth are claw-shaped.  Ctenitis paranaensisand  C. falciculataalso have dark brown to blackish scales on rachis, however, in these two species the scales are entire or slightly denticulate, and on pinnae abaxially the scales are only on costa, not on costule (but with or without proscales), while in  C. distansthere are scales on costa and costule (at least basally). We recognize two varieties of this species:  C. distansvar. distansand  C. distansvar. isabellina, as follows.  38• Phytotaxa335 (1) © 2018 Magnolia Press VIVEROS ET AL.A TAXONOMIC MONOGRAPH OF  CTENITISINSOUTH AMERICA   FIGURE 18.Distribution of four taxa of  CtenitisinSouth America. A:  C. distansvar. isabellina. B:  C. equestrisvar. equestris. C:  C. eriocaulis. D:  C. falciculata.  Phytotaxa335 (1) © 2018 Magnolia Press • 39  8.1.  Ctenitis distans(Brack.) Ching (1938: 277)var. distans .Figs. 09G, 15A–C, 16D.      Aspidium ctenitis Link (1833: 122).  Nephrodium ctenitis(Link) Bakerin  Hooker & Baker (1868: 265).   Dryopteris ctenitis(Link) Kuntze (1891: 812). Type: — BRAZIL(from Brazil, cultivated in Hort. Berol.). Unknown s.n.( lectotypeB 20 0054401!, designated here, isolectotypesB 20 0054400! BM 000937853!).     Aspidium amaurolepis Fée (1869: 137).   Dryopteris ctenitis(Link) Kuntze f. amaurolepis(Fée) Christensen (1913a: 94). Type: — BRAZIL. Rio de Janeiro, Glaziou 2370( lectotypeP00170066!, designated here, isolectotypesB 20 0067955!, K 000590326!, P 00170068!); remaining syntypes: — BRAZIL. Rio de Janeiro, Glaziou 1680(P 02141712!, P 00170065!, US00067052!); Glaziou 2371(P 00170067!, not P 00170069!, P 02141714!); Glaziou 2372(BR 0000013531621!, P 00170070!, P 00170072!, S 14-32612!, not K 000590327!, P 00170071!, P 01567696!).   Selected specimens examined:—  BRAZIL. Alagoas: São José da Lage, Usina Serra Grande, Mata Maria Maior, Mata do Pinto(área do Projeto Capivara),  380–507 m, 08º58’19.01”S, 36º06’36.05”W,  25 November 2001,  Pietrobom5392( BHCB);  Bahia: Amargosa, Área de Duas Barras,  625 m, 14°09'66" S, 39°49'44" W  19 October 2007,  Paixão& Nascimento1412( HUEFS);  Camacan, RPPN Serra Bonita,  700 m, 15º23’35”S, 39º33’53”W,  14 April 2007,  Matoset al. 1372( NY);  Santa Terezinha, Serra da Jibóia,  3 March 2001,  Nonatoet al. 800( HUEFS);  Ceará: Baturité, Sítio Santa Clara, 900,  9 December 1937,  Eugenio46( RB);  Espírito Santo: Santa Teresa, Reserva Biológica Augusto Ruschi, Trilha da Preguiça,  830–900 m, 19º54’58”S, 40º32’31”W,  3 December 2008,  Salinoet al. 14055( BHCB);  Minas Gerais: Nova Lima, RPPN Matado Jambreiro,  770 m, 19º58’41.9”S, 43º53’10.6”W,  17 September 2003,  Figueiredoet al. 45( BHCB);  Viçosa, Fazenda Faziuma,  700 m,  27 May 1930,  Mexia4750(F, GH, MO, NY, UC);  Paraná: Telêmaco Borba, Parque Ecológico da Klabin, Trilha Ecológica,  850 m, 24º12’42”S, 50º33’26”W,  1 February 2005,  Sakagami337 et al.( BHCB); Pernambuco:  Cabo de SantoAgostinho, Reserva de Gurjaú,  14 March 1991,  Fonsecaet al. s.n.( MO);  Jaqueira, Usina Colônia,  652 m, 8°43'00" S, 35°50'20" W,  31 May 2001,  Lopes& Pietrobom277( RB);  Timbaúba, Complexo da Serra do Mascarenhas, Usina Cruangi, Engenho Água Azul, ca.  304–394 m, 07º36’31.5”S, 35º22’42.9”W,  13 November 2000,  Pietrobomet al. 4650( BHCB); Rio de Janeiro:  Rio de Janeiro, Tijuca,  500–975 m,  31 March 1929,  Smith& Brade2235( GH);  Petrópolis, próximo à Carangola,  910 m, 22º27’53”S, 43º10’02”W,  22 May 2010,  Heringer469( BHCB);  São Paulo: Analândia, Salto do Rio Corumbataí,  25 June 1993,  Salino1791( BHCB).  Habitat and distribution:—Terrestrial, endemic to Atlantic Forest, 200–1200 m. Northeastern to southern Brazil( Fig. 16D; Tab. 01).   Notes:—In this typical variety thescales on costa abaxially are castaneous, but dark brown to blackish towards apex, distinctly clathrate, lanceolate, to 0.7 mmwide; catenate trichomes are absent on abaxial pinnae axes and laminar surface between veins glabrous or with sparse bacilliform trichomes ( Figs. 15A –C). In opposition,  C. distansvar. isabellinascales that are on costa abaxially are quite uniformly dark brown to blackish, subclathrate, linear to linear-lanceolate, to 0.3 mmwide ( Fig 15E); catenate trichomes are sparse on abaxial pinnae axes, as well on laminar surface between veins. Specimens of  C. distansvar. distanshave 4 vascular bundles at petiole base (Fig. 02B), while  C. distansvar. isabellinahave 6 (Fig. 02D). The vascular bundles at petiole base are an interesting feature to be more investigated in  Ctenitis, once the number of them is not constant under the current genus circumscription and even not constant in some of the species.  Fée (1869)and Christensen (1913a)were aware of the distinction of those two taxa. Fée (1869)considered them as species (  Aspidium amaurolepisand  A. isabellinum) and Christensen (1913a)as form undera single species (  Dryopteris ctenitisf. amaurolepisand  D. ctenitisf. isabellina). Brade (1972)identified several herbarium sheets with these infraspecific names, although he has not recognized them in his treatment for  Dryopterisin Brazil. Notwithstanding, after analyzing the typesand several materials, we also recognize these taxa and, despite the differences we were able to diagnose, there is much overlap of characters between them, that seems unwise recognizing them as different species at this moment.  Link (1833), when describing  Aspidium ctenitis, did not cite any specimen. Christensen (1913a)cited that the type was in B. Indeed, in B there are two sheets that certainly correspond to original material, once they are labeled with such name and with “H.B. Link”. Probably they are from a single specimen, but this is not clearly labeled. Then, we chose as lectotype(Art. 8.3, Rec. 8A.4, Art. 9.2, 9.5, 9.11, 9.12, 40 Note 1 of the ICN— McNeill et al. 2012) the sheet with more pinnae. A fragment removed from one of these sheets is in BM (purchased from Christensen’s herbarium). The plant from which the collection was made, was being cultivated in the Horto Berolinense, catalogued as from Brazil. As mentioned by Christensen (1913a)it seems to be an abnormal form. Its  40• Phytotaxa335 (1) © 2018 Magnolia Press VIVEROS ET AL. basal pinna is more developed, pinnate at base (with pinnules proximally) and with segments (distally) serrate. Nevertheless, the other features are the same as the typematerial of  Lastrea distansand most specimens of  Aspidium amaurolepis. The names  Aspidium amaurolepisand  A. isabellinumare based on Glaziou’scollections. For  A. amaurolepis, Fée (1869)cited Glaziou 1680, 2370, 2371and 2372and for  A. isabellinumonly Glaziou 2368. By Fée’s original descriptions,  A. amaurolepismatches with the same taxon of  A. ctenitisand  L. distans, here recognized as the typical variety of  C. distans, while  A. isabellinum,corresponds to  C. distansvar. isabellina(here combined). However, there are several sheets of Glaziou’s collections and some of them do not correspond to the taxon they supposed to be (see the headers 8.1. and 8.2.). Following the Art. 8.2, 8.3, 9.11, 9.12 and 9.14 of ICN( McNeill et al.2012), we propose a lectotypefor each name. For  A. amaurolepisthe sheet chosen is the one with entire petiole and a large portion of lamina, corresponding to  C. distansvar. distans. For  A. isabellinum, we chose the sheet that corresponds to  C. distansvar. isabellinaand is with the original Fée’s label with his signature.  8.2.  Ctenitis distans(Brack.) Ching var. isabellina(Fée) R.S. Viveros & Salino,  comb. nov., stat. nov.Figs. 09H, 15D–E, 18A.  Aspidium isabellinum Fée (1869: 137).  Dryopteris isabellina(Fée) Christensen (1906: 272).  Dryopteris ctenitis(Link.) Kuntze f. isabellina(Fée) Christensen (1913a: 94). Type:— BRAZIL. Rio de Janeiro, Glaziou 2368( lectotypeP 00170017!, designated here, isolectotypesB 20 0038358!, BM 000937854!, P 00643495!, not B 20 0054404!, K 000880597!, P 00170018!, P 01630476!, S 06-528!).   Selected specimens examined:—  BRAZIL. Espírito Santo: Estradado  Espírito Santo,  10 November 1946,  Bradeet al. 8072( RB);  Minas Gerais: Carangola, Fazenda Boa Vista,  30 May 1989,  Salinoet al. 820( UEC);  Caratinga, Estaçãoecológica de Caratinga( Fazenda Montes Claros),  6 September 1998,  Salino4323( BHCB);  São Sebastião do Paraíso, Fazenda Calada,  16 April 1945,  Brade& Barbosa17956( RB);  Paraná: Jaguariaiva, Serrinha,  18 November 1976,  Hatschbach39235( MBM, UC); Rio de Janeiro:  Rio de Janeiro, Parque Nacional da Tijuca,  17 June 1967,  Suthers88( UC);  Santa Maria Madalena,  1200 m  4 March 1934,  Lima& Brade13137( RB);  São Paulo: Analândia, Serra do Cuscuzeiro,  25 October 1986,  Salino32a( UEC);  Itapetinga,  26 June 1914,  Brade7598( UC);  São Paulo, Reserva Florestal"Armando de Salles" USP,  12 September 1996,  Groppo Júnior217( NY);  Tapiraí, Reserva Votorantin, Complexo Juquiá, Trilha Boatudo,  425 m, 24º02’14”S, 47º22’53”W,  Salinoet al.15640( BHCB).  Habitat and distribution:—Terrestrial. Endemic to Atlantic Forest, 150–1100 m. Southeastern to southern Brazil( Fig. 18A; Tab. 01).   Notes:—This variety differsfrom the typical by dark brown to blackish among some castaneous scales on costa abaxially, which are linear or linear-lanceolate, to 0.3 mmwide and sparse catenate trichomes on costa, costule, veins and between veins abaxially ( Figs. 15D –E). See the notes about  C. distansvar. distansfor further information about morphology, differences between the varieties and typification. 2001-11-25 BHCB Projeto Capivara & Pietrobom Brazil Sao Jose da Lage 443 -8.971947 Mata Maria Maior 1 -36.110012 Usina Serra Grande 39 1 Alagoas 2007-10-19 HUEFS Paixao & Nascimento Brazil 625 Area de Duas Barras -39.828888 Amargosa 39 1 Bahia 2007-04-14 NY Matos Brazil 700 -15.393055 Serra Bonita 21 -39.56472 Camacan 39 1 [568,1432,679,704] 2001-03-03 HUEFS Nonato Brazil Serra da Jiboia Santa Terezinha 39 1 [151,1004,716,741] 1937-12-09 RB Eugenio Brazil Sitio Santa Clara Baturite 39 1 Ceara 2008-12-03 BHCB Salino Brazil Santa Teresa 865 -19.916111 Trilha da Preguica 21 -40.541943 Reserva Biologica Augusto Ruschi 39 1 Espirito Santo 2003-09-17 BHCB Figueiredo Brazil 770 -19.978306 Jambreiro 1 -43.88628 Nova 39 1 Minas Gerais 1930-05-27 Brazil 700 Fazenda Faziuma Vicosa 39 1 2005-02-01 BHCB Sakagami Brazil Telemaco Borba 850 -24.211668 Trilha Ecologica 21 -50.55722 Parque Ecologico da Klabin 39 1 Parana 1991-03-14 MO de Gurjau & Fonseca Brazil Cabo de Santo 39 1 Cabo de Santo 2001-05-31 RB Lopes & Pietrobom Brazil 652 -8.716666 Usina Colonia 21 -35.838886 Jaqueira 39 1 2000-11-13 BHCB Pietrobom Brazil Timbauba 349 -7.60875 Usina Cruangi 1 -35.378582 Complexo da Serra do Mascarenhas 39 1 [464,1431,1039,1064] 1929-03-31 GH Smith & Brade Brazil 738 Tijuca 39 1 Rio de Janeiro [151,1379,1076,1101] 2010-05-22 BHCB Heringer Brazil 910 -22.464724 Carangola 21 -43.167225 Petropolis 39 1 1993-06-25 BHCB Salino Brazil Salto do Rio Corumbatai Analandia 39 1 Sao Paulo [535,950,836,861] Estrada Brazil Espirito Santo 40 1 Espirito Santo 1946-11-10 RB Brade Brazil Espirito Santo 40 1 Espirito Santo [350,1308,871,896] 1989-05-30 UEC Salino Brazil Fazenda Boa Vista Carangola 40 1 Minas Gerais 1998-09-06 BHCB Salino Brazil Caratinga Caratinga Estacao 40 1 1945-04-16 RB Brade & Barbosa Brazil Fazenda Calada Sao Sebastiao do Paraiso 40 1 1976-11-18 MBM Hatschbach Brazil Serrinha Jaguariaiva 40 1 Parana 1967-06-17 UC Suthers Brazil Parque Nacional da Tijuca 40 1 Rio de Janeiro [475,1305,1015,1040] 1934-03-04 RB Brade Brazil 1200 Santa Maria Madalena 40 1 1986-10-25 UEC Salino Brazil Serra do Cuscuzeiro Analandia 40 1 Sao Paulo [942,1431,1052,1077] 1914-06-26 UC Brade Brazil Itapetinga 40 1 [151,1335,1087,1112] 1996-09-12 NY Groppo Junior Brazil Reserva Florestal 40 1 Sao Paulo BHCB Salino Brazil Tapirai 425 -24.037222 Complexo Juquia 21 -47.381386 Reserva Votorantin 40 1