Ctenitis refulgens (Klotzsch ex Mett.) Christensen ex Vareschi (1968: 404)

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

22. Ctenitis refulgens (Klotzsch ex Mett.) Christensen ex Vareschi (1968: 404) . Phegopteris refulgens Klotzsch ex Mettenius (1864: 240) . Polypodium refulgens (Klotzsch ex Mett.) Hooker & Baker (1868: 307) . Nephrodium refulgens (Klotzsch ex Mett.) Diels (1899: 170) . Dryopteris refulgens (Klotzsch ex Mett.) Christensen (1906: 288) . Type:— GUYANA. Schomburgk 1183 (lectotype B 20 0064265!, designated here, isolectotype B 20 0064260!); remaining syntypes:— COLOMBIA. Chocó (New Granada), Schott 07 ( US 01105867!, US 01105866!); Magdalena, 100 m, Lindig 382 (B 20 0064261!); Tolima: Monte Tolima, Linden 1011 (not found).

Stems ascending or short-creeping, 1.7–3.0 cm diam., scales 4.0–11.1 × 0.2–0.6 mm, light castaneous, subclathrate, linear, entire, without fimbriae; leaves 48.5–121 cm long; petioles 15–54 cm × 2.0–6.0 mm, with 4 vascular bundles at base, stramineous or tan, scales 2.4–10.0 × 0.1–0.6 mm, light castaneous, subclathrate, tangled on petiole base, becoming patent or ascending towards distal portion, flattish, flaccid, linear with truncate base and filiform apex, entire, without fimbriae, sparse catenate trichomes abaxially, sparse glandular trichomes; laminae 33.5–67 × 21.5–35 cm, width 1/2–2/3 of length, 1-pinnate-pinnatifid basally, medially and apically, lanceolate or ovate, apex confluent; rachises stramineous or tan, scales like those on distal portion of petioles, sparse catenate trichomes abaxially, sparse glandular trichomes; pinnae 7–15 pairs, the basal and medial ones stalked to 3.8 mm long or sessile, the apical ones sessile, basal pinnae basiscopically and acroscopically somewhat equally developed, the medial 12.2–21.5 × 2.9–3.5 cm, lanceolate, incised ca. 1/2 (rarely to 2/3) of the distance between the segment apex and costa, basal segments as long or shorter than the next, apex attenuate; adaxial pinnae axes with sparse scales on costa, 1.8–3.1 × 0.06 mm, castaneous, filiform, catenate trichomes dense on costa, bacilliform trichomes absent; adaxial laminar surface between veins glabrous; abaxial pinnae axes with sparse scales on costa, 1.5–2.5 × 0.1–0.2 mm, castaneous or light castaneous, clathrate, patent or ascending, flattish or twisted, flaccid, linear with truncate or slightly cordate base and filiform apex, entire, with or without some short fimbriae at base and laterally, proscales absent, catenate trichomes absent or rare on costule, bacilliform trichomes sparse on costa, costule and veins, glandular trichomes sparse on costa, filiform trichomes absent; abaxial laminar surface between veins with sparse bacilliform trichomes; segments 16–23 pairs, 6.2–9.1 mm wide, patent or subfalcate, entire or repand, apex

A TAXONOMIC MONOGRAPH OF CTENITIS IN SOUTH AMERICA

Phytotaxa 335 (1) © 2018 Magnolia Press • 65 obtuse, margin with catenate trichomes, the distance from each other is narrower than segments width; veins simple, 7–12 pairs per segment, the basal ones from adjacent segments reach the margin at sinus, sometimes one of these veins ends before the margin towards the sinus, the next one or two pairs of basal veins reach the margin somewhat above the sinus; sori medial or inframedial, indusia absent; spores with coarse folds and large tubercles.

Notes:— Christensen (1913a) proposed the recognition of two infraspecific taxa under C. refulgens (treated by him as Dryopteris refulgens ). According to his descriptions it seemed that the variety peruviana were just smaller and younger plants from Peru. However, after analyzing numerous exsiccatae, it seems reasonable to recognize the two taxa, once larger specimens from Peru and Bolivia showed the same characters indicated by Christensen (1913a) to D. refulgens var. peruviana .

66 • Phytotaxa 335 (1) © 2018 Magnolia Press

VIVEROS ET AL.

Mettenius (1864) cited four collections in the protologue of Phegopteris refulgens : Linden 1011 (from Tolima, Colombia), Lindig 382 (from Magdalena, Colombia), Schott 7 (from Panama, currently Colombia in Chocó) and Schomburgk 1183 (from Guyana). Christensen (1913a) erroneously cited as type the Schomburgk 1128 (not cited in the protologue) and Schomburgk 1183 (cited in the protologue). Lellinger (1977) did not consider or corrected the typification made by Christensen (1913a), and chose as lectotype Schott 7 (two sheets in US), presuming that an isolectotype should be in B. Nonetheless, Tryon & Stolze (1991) and Moran (1995) attributed the lectotypification to Christensen (1913a), understanding that the lectotype should be Schomburgk 1183 in B. Mickel & Smith (2004), however, gave credit to the lectotype chosen by Lellinger (1977), also presuming that a duplicate should be in B. In our visits to B, we did not find any Schott 7. According to Art. 7.10 and 9.19 of ICN ( McNeill et al. 2012), the first typification proposed by Christensen has priority, as presumed by Tryon & Stolze (1991) and Moran (1995). Therefore, Schomburgk 1183 in B must be considered as the lectotype (Art. 9.9 of ICN — McNeill et al. 2012), since the other collection cited by Christensen (1913a) was not cited in the protologue. There are, however, two sheets of that gathering in B. Both sheets may correspond to the same individual, but there is no label that clearly supports this hypothesis. Therefore, the two sheets must be considered as duplicates (Art. 8.3 of ICN — McNeill et al. 2012). This means that Christensen (1913a) conducted the first step lectotypification by citing the gathering Schomburgk 1183. Here we conduct the second-step lectotypification by designating the sheet B 20 0064265! as the lectotype.

22.1. Ctenitis refulgens (Klotzsch ex Mett.) C. Chr. ex Vareschi (1968: 404) var. refulgens . Figs. 05B, 10I, 24A–C, 25C.

Phegopteris tricholepis Fée (1869: 98) . Type : — BRAZIL. Amazonas : Prope San Gabriel da Cachoeira , ad Rio Negro, brasilie borealis, Spruce 2100 (lectotype P 00170060!, designated here, isolectotypes BR [AWH collection], K 000954421!, P 00170064!, P 01449837!, RB 00657603!).

Selected specimens examined:— BOLIVIA. Beni: Gral. Ballivián, ca. 33 km south of San Borja , along a logging track, 264 m, 15º09’S, 66º44’W, 27 June 2009, Weigelt et al. 90254 ( UC) ; La Paz: Nord Yungas, Polo–Polo bei Coroico , 1100 m, October–November 1912, Buchtien 3409 (F) ; Pando: Manuripi, camino Cobja-Chivé, comunidad Holanda , entrando por el camino al puesto militar adelantando Dolores , antes de cruzar el rio Manurimi , 230 m, 11º47’S, 68º42’W, 28 July 2003, Jimenez 1954 ( UC) ; BRAZIL. Acre: Cruzeiro do Sul, comunidade de Santa Luzia , 275 m, 07º53’45’’S, 72º24’30’’W, 10 December 2010, Salino & Almeida 15004 ( BHCB) ; Porto Walter, Igarapé Ouro Preto , 20 November 2001, Croat 85538 ( RB) ; Amazonas: Prope San Gabriel da Cachoeira, ad Rio Negro , brasilie borealis, Spruce 2100 ( RB) ; Tefé , northwest shore, 3º20’S, 64º50’W, 11-14 December 1982, Plowman et al. 12497 ( INPA) ; Mato Grosso: Alta Floresta, Assentamento Rural de Carlinda , 10°00' S, 56°00' W, 6 May 1986, Windisch 4755 ( UC) ; Aripuanã, Dardanelos , 28 September 1975, Lisboa et al. 405 ( INPA) ; Paranaíta, Propriedade do Sr. Nilo Weber , 9°20'34" S, 56°46'15" W, 18 July 2008, Dias-Melo et al. 584 ( RB) ; Pará: Canaã dos Carajás, Serra do Tarzan , 15 December 2012, Salino 15575 ( BHCB) ; Marabá, Serra Norte , 20 May 1982, Secco et al. 291 ( RB) ; Novo Repartimento, Zona de Preservação da Vida Silvestre, Base 3, 10 July 2008, Fernandes 167 ( RB) ; W bank of Rio Maicuru , 0°55' S, 54°26' W, 29 July 1981, Strudwick et al. 3718 (F) ; Roraima: Caracaraí, Parque Nacional do Viruá , 01º29’25’’S, 61º00’38’’W, 13 October 2011, Mota et al. 2351 ( BHCB) ; COLOMBIA. Chocó: Río Sucio, Parque Nacional Natural Los Katios , 290 m, 29 May 1976, Forero et al. 1646 ( MO) ; ECUADOR. Napo: Yasuní National Park , 200–300 m, 00 o 42’S, 76 o 29’W, 18 April 1997, H. Tuomisto et al. 10676 ( UC) ; Tena, Jatun Sacha Biological Station , 400 m, 1°04' S, 77°36' W, 9 December 1990, Fay & Fay 2678 ( UC) ; FRENCH GUIANA. Monts la Fumée , 200–400 m, 3°37' N, 53°12' W, 2 September 1982, Boom & Mori 1588 ( NY) ; Pentes exposées au nord, à 300 m au nord du camp Guyanor, à 1,3 Km de St Elie ( Guyane ), 4°49' N, 53°18' W, 31 July 1997, Boudrie 2838 ( UC) ; Roche Koutou–Bassin du Haut–Marouini , 165 m, 02 o 52’S, 54 o 04’W, 17 August 1987, Granville et al. 9420 ( UC) ; GUYANA. Demerara, Berbier River , 1897, G. S. Jenman s.n. ( NY) ; PERU: Madre de Dios: Tambopata, Puerto Maldonado , 12°64' S, 69°33' W, 30 January 2006, Chambi & Chambi 178 ( NY) ; Manu, 10 km E from the mouth of Río Manu , northern bank of Río Madre de Díos, 300–400 m, 12º15’S, 70º48’W, 25 October 1998, Tuomisto et al. 13397 ( UC) ; SURINAME. Sipaliwini, Central Suriname Nature Reserve, vicinity of camp at southern base of the first peak in Eilerts de Haan mountain range ca. 6 km ENE of Kayserberg Airstrip, 250–350 m, 03 o 09’00”N, 56 o 27’30”W, 23 June 2003, Herrera & Koemar 10044 ( UC) ;

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Phytotaxa 335 (1) © 2018 Magnolia Press • 67 VENEZUELA. Apure: Paez , 25 km E of El Nula, 200 m, 07 o 14’N, 71 o 45’W, 1 July 1983, van der Werff & Gonzáles 4678 ( MO) ; Barinas: Pedraza, above El Agarrobo, 400m, 08 o 31’N, 70 o 35’W, 3 August 1983, van der Werff & Ortiz 5808 ( MO) ; Bolívar: Raul Leoni, 86 km al Este de San Francisco , 430 m, 06 o 09’N, 64 o 24’W, June1989, Delgado 254 ( MO) ; Island Margarita , 8 February 1901, Miller 169 ( MO) ; Zulia: Bolívar, Cuenca del Embalse Burro Negro ( Pueblo Viejo ), 250–300 m, 5–8 August 1980, G. S. Bunting 9577 ( UC) .

Habitat and distribution:— Terrestrial in rainforests, 100–900 m. Mesoamerica ( Mexico, Belize, Guatemala, Honduras, Nicaragua, and Panama), West Indies (Trinidad) and South America (French Guiana, Suriname, Guyana, Venezuela, Colombia, Ecuador, Peru, Bolivia and Brazil; Fig. 25C ; Tab. 01) .

Notes:—The typical variety is recognized by pinnae incised 1/2–2/3 of the distance between segment apex and costa (Fig. 05B–C), and light castaneous and linear scales on costa abaxially. Ctenitis refulgens var. refulgens is similar to C. nervata and C. aspidioides (see corresponding notes) by pinnae incision, differing from both by light castaneous, which resemble C. eriocaulis , however not subulate, but linear ( Figs. 24A–C ). Furthermore, C. eriocaulis pinnae are incised more than 3/4, the indusia is present and the veins from adjacent segments reach the margin well above the sinus. In C. refulgens there are scales also on costa adaxially, while, in C. nervata and C. aspidioides there are not. Furthermore, one vein from adjacent segments can sometimes ends before margin towards the sinus, or the next one or two pairs reach the margin somewhat above the sinus.

Fée (1869) cited the collection Spruce 2100 for Phegopteris tricholepis . No holotype or herbarium were specified. Sheets of this collection were found in P, BR, K, and RB. Supported by the Art. 8.3, Rec. 8A. 4, Art. 9.2, 9.5, 9.11, 9.12, 40 Note 1 of ICN ( McNeill et al. 2012), we designate here for lectotype one of the sheets in P, which contains the most complete leaf sample.

22.2. Ctenitis refulgens (Klotzsch ex Mett.) C. Chr. ex Vareschi var. peruviana (C. Chr.) R.S. Viveros & Salino , comb. nov. Figs. 05A, 10J, 24D–E, 25D. Dryopteris refulgens (Klotzsch ex Mett.) Christensen (1906: 288) var. peruviana Christensen (1913a: 90) . Type:— PERU. Prope Tabalosas inter urbem Moyobamba et fluvium Río Huallaga, Stübel 1097 (lectotype B 20 0064268!, designated here); remaining syntypes:— PERU. Tarapoto, in Monte Guayrapurina, Spruce 4712 ( US 00513182!, isosyntype MO 255866!); PERU. San Martín: Tarapoto, in Monte Campaña, Spruce 4657 (LD 1765418, B 20 0064269!, isosyntypes BM 000777157!, BM 000777156!, BR 0000013514594!, GH 00443892!, K 000200138!, K 000200139!, K!, P 00642693!).

Selected specimens examined:— BOLIVIA. La Paz: Sud Yungas, Alto Beni, Sapecho, Colonia Tarapaca , 610 m, 15 o 32’S, 67 o 21’W, 28 Octuber 1997, Krömer et al. 50 ( UC) ; PERU. Huanuco: Pachitea, Puerto Inca , 350 m, 09 o 31’S, 74 o 58’W, 13 April 1982, Smith 1299 ( MO) ; Madre de Díos: Tambopata, Las Piedras. Cusco Amazónico , 200 m, 12°29' S, 69°03' W, 11 November 1991, Timaná & Jaramillo 3136 ( MO) ; Manu, close to the Village of Diamante, Southern side of Río Alto Madre de Díos, 400 m, 12 o 20’S, 70 o 57’W, 24 October 1998, Tuomisto et al. 13392 ( UC) ; San Martín: Lamas, Alonso Alvarado, San Juan de Pacayzapa , al leste del Puente (carretera a Moyobamba), 16 April 1973, Schunke 5941 (F) .

Habitat and distribution:—Terrestrial in rainforests, 230–1180 m. South America ( Peru and Bolivia; Fig. 26D ; Tab. 01).

Notes:— Ctenitis refulgens var. peruviana differs from the typical by the pinnae incision 1/4–1/3 of the distance between the segment apex and costa (Fig. 05A), and by darker (castaneous) and lanceolate scales on costa abaxially ( Fig. 24D–E ), once the typical is incised 1/2–2/3 and the costal scales abaxially are lighter (light castaneous) and linear.

When proposing this variety, Christensen (1913a) cited four syntypes: Spruce 4657 (B, LD), Spruce 4712 ( US), Stübel 1097 (B). This last we designate as lectotype (Art. 9.12 of ICN— McNeill et al. 2012).

Figures

FIGURE 24. Some morphological characters of Ctenitis from South America. A–C: Ctenitis refulgens var. refulgens (Salino 15242, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: abaxial surface of rachis, costa and segment, showing indument. C: abaxial costa scale. D–E: C. refulgens var. peruviana (Tuomisto, 13392 UC). D: abaxial surface of a portion of medial pinna, showing indument. E: abaxial costa scale. Scale bars in A = 1 cm, B–E = 1mm.

FIGURE 25. Distribution of four taxa of Ctenitis in South America. A: C. nigrovenia. B: C. paranaensis. C: C. refulgens var. refulgens. D: C. refulgens var. peruviana.

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.