Ctenitis nigrovenia (Christ) Copeland (1947: 124)

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

20. Ctenitis nigrovenia (Christ) Copeland (1947: 124) . Figs. 02A, 10G, 22C–D, 25A. Nephrodium nigrovenium Christ in Donnell Smith (1895: 545). Dryopteris nigrovenia (Christ) Christensen (1906: 279) . Type:— HONDURAS. Santa Bárbara: San Pedro Sula, September 1887, 1000 ft., Thieme s.n. [Herb. Donnell Smith 5646] (lectotype P 00642702! designated by Tryon & Stolze 1991, isolectotype US 00067035!).

Aspidium setosum Klotzsch (1847: 371) , nom. illeg., non ( Thunberg 1784: 337) Swartz (1801: 39), non Wallich (1828: 371), nom. nud. Type: — COLOMBIA. Moritz 204 (lectotype P 01514077!, designated here, isolectotypes LE!, P 04021631!, P!, US 00067067!).

Aspidium tonduzii Christ (1901: 34) . Dryopteris tonduzii (Christ) Christensen (1906: 664) . Ctenitis tonduzii (Christ) Tryon & Tryon (1982a: 215) . Type: — COSTA RICA: Forêt de Tuis, Tonduz 11333 (lectotype P 00630760!, designated here, isolectotypes!, BM 000904585!, MO 122617!, P 00642673!, U 0007377!, US 00067047!, US 1100787!).

Ctenitis thelypteroides Smith (1975: 215) . Type: — MEXICO. Chiapas: Las Margaritas, western side of Laguna Miramar , E of San Quintín , Breedlove 33280 (holotype CAS 0001586!, isotypes CAS 0001585!, MEXU 00301602 [image!], NY 00179358!).

Stems erect, ascending or short-creeping, 0.9–3.3 cm diam., scales 2.1–8.0 × 0.4–1.5 mm, castaneous, subclathrate or clathrate, lanceolate, entire, without fimbriae; leaves 32–100 cm long; petioles 12–50 cm × 1.2–2.3 mm, with 3 vascular bundles at base, stramineous, scales 2.4–6.8 × 0.3–1.2 mm, castaneous on petiole base, becoming dark brown to blackish on distal portion, subclathrate or clathrate, tangled on petiole base, becoming patent or ascending towards distal portion, flattish, flaccid, lanceolate with truncate base and filiform apex, entire, with or without some short fimbriae at base, sparse catenate trichomes abaxially, sparse glandular trichomes; laminae 20–50 × 9.7–25.5 cm, width ca. 1/2 of length or somewhat wider, 1-pinnate-pinnatisect basally, 1-pinnate-pinnatisect or 1-pinnate-pinnatifid, almost pinnatisect medially and apically, lanceolate or ovate, apex confluent; rachises stramineous, scales like those on distal portion of petioles, sparse catenate trichomes abaxially, sparse glandular trichomes; pinnae 7–16 pairs, the basal and medial ones stalked to 4.5 mm long, the apical ones sessile, basal pinnae basiscopically and acroscopically somewhat equally developed, the medial (5) 7–15 × 1.4–3 cm, lanceolate, incised more than 3/4 of the distance between the segment apex and costa, basal segments somewhat shorter than the next at basal pinnae, but longer at medial, apex attenuate; adaxial pinnae axes scales absent, catenate trichomes, dense on costa, sparse on costule, rare on veins, bacilliform trichomes absent; adaxial laminar surface between veins glabrous; abaxial pinnae axes with sparse scales on costa, 0.6–2.2 × 0.2–0.3 mm, dark brown, clathrate, ascending, mostly flattish, but can be vaulted at base, flaccid, linear-lanceolate with truncate or slightly cordate base and filiform apex, entire, with or without some short fimbriae at base, proscales absent, catenate trichomes sparse on costa, costule and rare on veins, bacilliform trichomes sparse on costa, costule and veins, glandular trichomes absent, filiform trichomes absent; abaxial laminar surface between veins with sparse bacilliform trichomes; segments 13–19 pairs, 2.8–5.7 mm wide, patent or subfalcate, serrate towards apex, rarely entire, apex acute or apiculate, margin glabrous, the distance from each other is narrower than segments width; veins simple or 1-forked, 6–11 pairs per segment, the basal ones from adjacent segments end at margin well above the sinus; sori medial, inframedial or supramedial, indusia conspicuous or inconspicuous, entire, with bacilliform trichomes; spores coarsely echinate.

Selected specimens examined:— BOLIVIA. La Paz: Sud Yungas, Colonia Buena Vista, 11 August 1994, Seidel & Vaquita, D. 7622 ( UC); Santa Cruz: Florida, Refúgio Los Volcanes , 1050 m, 18°06' S, 63°36' W , 3 October 1997, Kessler et al. 12268 ( UC); Velasco, Parque Nacional Noel Kempff Mercado , 700 m, 14°31'16" S ,

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Phytotaxa 335 (1) © 2018 Magnolia Press • 61 60°44'14" W, 4 July 1996, Peña-Chocarro et al. 69 ( NY); BRAZIL. Mato Grosso: Serra Ricardo Franco , 500 m, 15ºS, 60ºW, 15 December 1977, Windisch 1498 ( UC) ; Pará: Canaã dos Carajás, Serra do Tarzan , 15 December 2012, Salino 15575 ( BHCB) ; COLOMBIA. Magdalena: Santa Marta, 1898, Smith 2581 (MO, NY) ; Risaralda: Pereira, Hacienda Los Visos , 11 August 1991, Silverstone-Sopkin & Arroyo 6295 (MO, UC) ; PERU. Cusco: La Convención, Rio Manguriari , 2 February 1991, Nuñez & Ortiz 12760 ( MO) ; San Martin: Hara, near Moyobamba , 04 June 1947, Woytkowski s.n. ( UC) ; VENEZUELA. Barinas: Bolívar, 600 m, 28 July 1984, Moran 3717 ( UC) ; Tovar , 1854, Fendler 194 ( MO) ; Lara: Jiménez, Parque Nacional Yacambú , 620–740 m, 9°41' N, 69°30' W, 24 October 1982, Davidse & Gonzáles 71020 ( MO) ; Portuguesa: Ospino , 900 m, 9°28'30" N, 69°28'40" W, 22 November 1990, González & Stergios 19 ( UC) ; Zulia: Lagunillas, Cuenca del Embalse Burro Negro ( Pueblo Viejo ), 550–600 m, 10°25' N, 70°49' W, 1 April 1982, Bunting et al. 11168 ( NY) .

Habitat and distribution:— Terrestrial, less frequently epipetric, in mountain forests, 50–900 m. Mesoamerica ( Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica and Panama), West Indies (Trinidad) and South America ( Venezuela, Colombia, Peru, Bolivia and Brazil; Fig. 25A ; Tab. 01) .

Notes:— Ctenitis nigrovenia is usually recognized by segments margin serrate and glabrous in a 1-pinnate-pinnatifid to 1-pinnate-pinnatisect lamina. Some specimens from Costa Rica and Mexico are smaller and because of that, the segments are repand or entire. These specimens have already been recognized as a distinct species, named C. tonduzii , but the microscopic characters are the same, for example bacilliform trichomes on costula, vein and between veins abaxially ( Fig. 22C ). Despite the absence of catenate trichomes on segments margin, the following further characters also contribute to recognized C. nigrovenia : scales on costa abaxially dark brown to blackish, indusia conspicuous with bacilliform trichomes, or inconspicuous and reduced, remaining a tuft of bacilliform trichomes. Ctenitis nigrovenia is like C. abyssi and C. laetevirens (see both for differences).

The collection cited in original description of Nephrodium nigrovenium is Thieme 5646 ( Donnell Smith 1895). As usual at that time, no holotype was designated, neither a herbarium was specified. One sheet of this collection is in P and another is in US. The one in P has a stamp of Christ’s Herbarium. Christensen (1913a) informed that the types were in Christ Herbarium and in US. The collections of Christ, the author of N. nigrovenium , were incorporated by the Roland Bonaparte’s herbarium, which was incorporated by P. Tryon & Stolze (1991) cited “P?” for the holotype and US for an isotype. Then, we consider this an inadvertent lectotypification and an error to be correct to lectotype (Art. 7.10, 9.9, 9.19 of ICN — McNeill et al. 2012).

Aspidium setosum ( Klotzsch 1847) is a later homonym, therefore illegitimate, but also susceptible to typification (Art. 7.5 of ICN — McNeill et al. 2012). Christensen (1913a) had already considered this name as a synonym of Dryopteris nigrovenia , and he attributed the collection Moritz 204 as from Tovar in Venezuela, not Colombia, as mentioned in the protologue. Moritz collected plants in Colombia and also in Venezuela, but the protologue and the herbarium sheets we found do not mention the locality “Tovar”. Although Christensen (1913a) has cited this collection in B, Christ’s Herbarium and S, only five sheets with the exact collecting number, written on them A. setosum , were found: three in P, one in LE and other with a pinna in US. The ones in P were incorporated from Luerssen-Weigel’s Herbarium. They are certainly original material (Art. 9.3 of ICN — McNeill et al. 2012), and here we designate as lectotype (Art. 8.3, Rec. 8A.4, 9.2, 9.5, 9.11, 9.12 and 40 Note 1 of ICN — McNeill et al. 2012) a specimen in P with a pretty leaf.

Christ (1901) cited the collection Tonduz 11333 for Aspidium tonduzii . Sheets of this were found in four herbaria. Supported by Art. 8.3, Rec. 8A.4, Art. 9.5, 9.12 and 40 Note 1 of ICN ( McNeill et al. 2012) we selected as lectotype one sheet in P, which is with Christ’s Herbarium stamp. Christensen considered A. tonduzii as Dryopteris tonduzii differing it from D. nigrovenia mainly by the lamina lighter and glabrous between veins on both surfaces. The types are almost glabrous, but they do contain the same indument of C. nigrovenia .

Ctenitis thelypteroides ( Smith 1975) were placed as synonym of C. nigrovenia by Stolze (1981). Moran (1995) said that C. nigrovenia was too variable and that more collections were need to clarify if C. thelypteroides and C. tonduzii could be distinct from C. nigrovenia . Mickel and Smith (2004), however, still maintained C. thelypteroides as distinct, separating by the presence of indusia in opposition to an exindusiate sori in C. nigrovenia . However, the types of C. nigrovenia ( N. nigrovenium ) are indusiate. The types of C. tonduzii and C. thelypteroides are quite similar in leaf size and segments margin, which are not so perfectly serrate as the typical C. nigrovenia . Furthermore, after analyzing several specimens from South to Mesoamerica and West Indies, we agree with Stolze (1981) and recognize all these three names as the same taxon. Therefore, C. nigrovenia is a widely distributed species with consequently morphological variations that such overlapping characters make unfeasible to distinguish more than one species.

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Christensen (1913a) presents the name Aspidium deltoideum Fournier (1872: 93) , non ( Swartz 1788: 133) Swartz (1801: 34), as a synomym of Dryopteris nigrovenia . However, Fournier did not intend to describe a new species, he just cited the name A. deltoideum of Swartz with the voucher Bourgeau 1644 from Mexico. It is an

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identification error of Fournier, such collection corresponds to C. nigrovenia and what Swartz (1801) referred is combined as Cyclosorus deltoideus (Sw.) Mazumdar & Mukhopadhyay (2013: 18) .

Figures

FIGURE 22. Some morphological characters of Ctenitis from South America. A–B: C. nervata (Almeida 1682, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: abaxial costa scale. C–D: C. nigrovenia (Salino 15212, BHCB). C: abaxial surface of a pinna, showing indument. D: abaxial costa scale. Scale bars in A = 1 cm; B = 0.5 mm; C, D = 1 mm.

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.