Ctenitis distans (Brack.) Ching (1938: 277)

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

8. Ctenitis distans (Brack.) Ching (1938: 277) . Lastrea distans Brackenridge (1854: 192) . Type:— BRAZIL. Rio de Janeiro, Capt. Wilkes s.n. (holotype US 00067060!).

Stems erect or ascending, 1.5–3.5 cm diam., 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 cm long; 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; rachises stramineous, scales like those on distal portion of petioles, rare to dense catenate trichomes abaxially, glandular trichomes absent; pinnae 10–37 pairs, the basal and medial ones stalked to 6.7 mm long, 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 axes with 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 veins glabrous; abaxial pinnae axes with 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 mm long 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 veins glabrous or with sparse bacilliform trichomes, or sparse catenate trichomes; segments 19–27 pairs, 2.8–5.6 mm wide, 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; veins simple, 7–15 pairs per segment, the basal ones from adjacent segments end at margin well above the sinus; sori inframedial, indusia conspicuous, entire or ciliate, with bacilliform trichomes; spores with coarse folds and large tubercles.

Notes:—The specimens of C. distans are 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 paranaensis and C. falciculata also 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. distans there are scales on costa and costule (at least basally). We recognize two varieties of this species: C. distans var. distans and C. distans var. isabellina , as follows.

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8.1. Ctenitis distans (Brack.) Ching (1938: 277) var. distans . Figs. 09G, 15A–C, 16D.

Aspidium ctenitis Link (1833: 122) . Nephrodium ctenitis (Link) Baker in Hooker & Baker (1868: 265). Dryopteris ctenitis (Link) Kuntze (1891: 812) . Type: — BRAZIL (from Brazil, cultivated in Hort. Berol.). Unknown s.n. (lectotype B 20 0054401!, designated here, isolectotypes B 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 (lectotype P00170066!, designated here, isolectotypes B 20 0067955!, K 000590326!, P 00170068!); remaining syntypes: — BRAZIL. Rio de Janeiro, Glaziou 1680 (P 02141712!, P 00170065!, US 00067052!); 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, Pietrobom 5392 ( BHCB) ; Bahia: Amargosa, Área de Duas Barras , 625 m, 14°09'66" S, 39°49'44" W 19 October 2007, Paixão & Nascimento 1412 ( HUEFS) ; Camacan, RPPN Serra Bonita , 700 m, 15º23’35”S, 39º33’53”W, 14 April 2007, Matos et al. 1372 ( NY) ; Santa Terezinha, Serra da Jibóia , 3 March 2001, Nonato et al. 800 ( HUEFS) ; Ceará: Baturité, Sítio Santa Clara , 900, 9 December 1937, Eugenio 46 ( RB) ; Espírito Santo: Santa Teresa, Reserva Biológica Augusto Ruschi, Trilha da Preguiça , 830–900 m, 19º54’58”S, 40º32’31”W, 3 December 2008, Salino et al. 14055 ( BHCB) ; Minas Gerais: Nova Lima, RPPN Mata do Jambreiro , 770 m, 19º58’41.9”S, 43º53’10.6”W, 17 September 2003, Figueiredo et al. 45 ( BHCB) ; Viçosa, Fazenda Faziuma , 700 m, 27 May 1930, Mexia 4750 (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, Sakagami 337 et al. ( BHCB) ; Pernambuco: Cabo de Santo Agostinho, Reserva de Gurjaú, 14 March 1991, Fonseca et al. s.n. ( MO) ; Jaqueira, Usina Colônia , 652 m, 8°43'00" S, 35°50'20" W, 31 May 2001, Lopes & Pietrobom 277 ( 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, Pietrobom et al. 4650 ( BHCB) ; Rio de Janeiro: Rio de Janeiro, Tijuca , 500–975 m, 31 March 1929, Smith & Brade 2235 ( GH) ; Petrópolis, próximo à Carangola , 910 m, 22º27’53”S, 43º10’02”W, 22 May 2010, Heringer 469 ( BHCB) ; São Paulo: Analândia, Salto do Rio Corumbataí , 25 June 1993, Salino 1791 ( 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 the scales on costa abaxially are castaneous, but dark brown to blackish towards apex, distinctly clathrate, lanceolate, to 0.7 mm wide; 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. distans var. isabellina scales that are on costa abaxially are quite uniformly dark brown to blackish, subclathrate, linear to linear-lanceolate, to 0.3 mm wide ( Fig 15E ); catenate trichomes are sparse on abaxial pinnae axes, as well on laminar surface between veins. Specimens of C. distans var. distans have 4 vascular bundles at petiole base (Fig. 02B), while C. distans var. isabellina have 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 amaurolepis and A. isabellinum ) and Christensen (1913a) as form under a single species ( Dryopteris ctenitis f. amaurolepis and D. ctenitis f. isabellina ). Brade (1972) identified several herbarium sheets with these infraspecific names, although he has not recognized them in his treatment for Dryopteris in Brazil. Notwithstanding, after analyzing the types and 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

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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 type material of Lastrea distans and most specimens of Aspidium amaurolepis .

The names Aspidium amaurolepis and A. isabellinum are based on Glaziou’s collections. For A. amaurolepis, Fée (1869) cited Glaziou 1680, 2370, 2371 and 2372 and for A. isabellinum only Glaziou 2368. By Fée’s original descriptions, A. amaurolepis matches with the same taxon of A. ctenitis and L. distans , here recognized as the typical variety of C. distans , while A. isabellinum , corresponds to C. distans var. 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 lectotype for each name. For A. amaurolepis the sheet chosen is the one with entire petiole and a large portion of lamina, corresponding to C. distans var. distans . For A. isabellinum , we chose the sheet that corresponds to C. distans var. isabellina and 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 (lectotype P 00170017!, designated here, isolectotypes B 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: Estrada do Espírito Santo, 10 November 1946, Brade et al. 8072 ( RB) ; Minas Gerais: Carangola, Fazenda Boa Vista , 30 May 1989, Salino et al. 820 ( UEC) ; Caratinga, Estação ecológica de Caratinga ( Fazenda Montes Claros ), 6 September 1998, Salino 4323 ( BHCB) ; São Sebastião do Paraíso, Fazenda Calada , 16 April 1945, Brade & Barbosa 17956 ( RB) ; Paraná: Jaguariaiva, Serrinha , 18 November 1976, Hatschbach 39235 ( MBM, UC) ; Rio de Janeiro: Rio de Janeiro, Parque Nacional da Tijuca , 17 June 1967, Suthers 88 ( UC) ; Santa Maria Madalena , 1200 m 4 March 1934, Lima & Brade 13137 ( RB) ; São Paulo: Analândia, Serra do Cuscuzeiro , 25 October 1986, Salino 32a ( UEC) ; Itapetinga , 26 June 1914, Brade 7598 ( UC) ; São Paulo, Reserva Florestal "Armando de Salles" USP, 12 September 1996, Groppo Júnior 217 ( NY) ; Tapiraí, Reserva Votorantin, Complexo Juquiá , Trilha Boatudo , 425 m, 24º02’14”S, 47º22’53”W, Salino et 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 differs from the typical by dark brown to blackish among some castaneous scales on costa abaxially, which are linear or linear-lanceolate, to 0.3 mm wide and sparse catenate trichomes on costa, costule, veins and between veins abaxially ( Figs. 15D – E ). See the notes about C. distans var. distans for further information about morphology, differences between the varieties and typification.

Figures

FIGURE 15. Some morphological characters of Ctenitis from South America. A–C: Ctenitis distans var. distans (Fontana 4996, BHCB). A: detail of abaxial surface of a medial pinna showing indument. B: abaxial costa scale. C: adaxial costa scale. D–E: C. distans var. isabellina (Lombardi 6830, BHCB). D: detail of abaxial surface of a medial pinna showing indument. E: abaxial costa scale. Scale bars = 1 mm.

FIGURE 16. Distribution of four taxa of Ctenitis in South America. A: C. bigarellae. B: C. christensenii. C: C. deflexa. D: C. distans var. distans.

FIGURE 18. Distribution of four taxa of Ctenitis in South America. A: C. distans var. isabellina. B: C. equestris var. equestris. C: C. eriocaulis. D: C. falciculata.