Ctenitis microchlaena (Fée) Stolze

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

18. Ctenitis microchlaena (Fée) Stolze in Tryon & Stolze (1991: 09). Figs. 01A, 10E, 21C–F, 23C. Aspidium microchlaena Fée (1857: 102) . Dryopteris microchlaena (Fée) Christensen (1906: 278) . Type:— MEXICO. Orizaba, Schaffner 459 (lectotype K 000590290! designated by Tryon & Stolze 1991).

Aspidium karstenii Braun (1858 : app. 3). Dryopteris karstenii (Braun) Christensen (1913a: 98) . Type:—“Unknown or unspecified Middle and South America”, probably VENEZUELA. February 1856, Gollmer s.n. (lectotype B 20 0058411!, designated here); remaining syntypes:— VENEZUELA. Karsten & Moritz s.n., (GH 00112619!, K 000590336!); COLOMBIA. Moritz 209 (BM 000937855!).

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Aspidium obtusilobum Fée (1857: 105) . Dryopteris huatuscensis Christensen (1906: 271) , non Dryopteris obtusiloba (Baker in Hooker & Baker 1868: 284) Kuntze (1891: 813), non ( Desvaux 1811: 357) Christensen (1906: 280), nom. illeg. Type: — MEXICO. Huatusco, Schaffner 213 (lectotype RB 00609505!, designated here).

Stems erect or ascending, 1.4–3.3 cm diam., scales 11.9–18.3 × (0.2) 0.6–1.2 mm, light castaneous or castaneous, sublclathrate, linear, entire, without fimbriae; leaves 72–123 cm long; petioles 24–47 cm × 2.5–3.1 mm, with 4 or 5 vascular bundles at base, brownish, tan or stramineous, scales 3.1–8.1 × 0.2–0.7 mm, castaneous, subclathrate, tangled on petiole base, becoming patent or ascending towards distal portion, flattish, flaccid, linear with truncate base and filiform apex, entire or slightly denticulate, with or without some short fimbriae at base, sparse to dense catenate trichomes abaxially, glandular trichomes absent; laminae 48–76 × 15.5–21.5 cm, width ca. 1/3 of length or narrower, 1-pinnate-pinnatifid basally, medially and apically, linear-lanceolate, apex confluent; rachises brownish, tan or stramineous, scales like those on distal portion of petioles, sparse to dense catenate trichomes abaxially, glandular trichomes absent; pinnae 19–27 pairs, the basal and medial ones stalked to 3.3 mm long, the apical ones sessile, basal pinnae basiscopically and acroscopically somewhat equally developed, the medial 6.5–13 × 1.3–2.2 cm, lanceolate, incised 2/3–3/4 or more of the distance between the segment apex and costa, basal segments as long or somewhat longer or shorter than the next, apex attenuate or acute; adaxial pinnae axes scales absent, catenate trichomes dense on costa, sparse on costule and veins, bacilliform trichomes absent; adaxial laminar surface between veins with sparse to dense catenate trichomes; abaxial pinnae axes with sparse scales on costa, 1.2–3.1 × 0.1–0.3 mm, castaneous, clathrate, ascending, flattish, flaccid, linear-lanceolate with truncate base and filiform apex, entire or slightly denticulate, with or without some short fimbriae at base, proscales to 0.9 mm long sparse on costule or absent, catenate trichomes sparse to dense on costa, costule and veins, bacilliform trichomes sparse on costa and costule or absent, glandular trichomes absent, filiform trichomes absent; abaxial laminar surface between veins with dense catenate and sparse bacilliform trichomes; segments 13–18 pairs, 3.3–5.7 mm wide, patent or subfalcate, entire or crenate, apex rounded, margin with catenate trichomes, the distance from each other is narrower than segments width; veins simple, 6–10 pairs per segment, the basal ones from adjacent segments end at margin well above the sinus; sori medial or supramedial, indusia conspicuous, entire, with bacilliform trichomes; spores with coarse folds.

Selected specimens examined:— BOLIVIA. La Paz: Pedro Domingo Murillo, valle de Zongo , 16 o 06’S, 68º07’W, 2600 m, 19 March 1995, Gonzales et al. 1893 ( UC) ; Santa Cruz: Andrés Ibañez , entre los 530–1121 m, 17º54’11”S – 17º53’42”S, 63º26’14”W – 63º28’17”W, 2 June 2007, Molina 215 ( MO) ; Velasco , Parque Nacional Noel Kempff M., 900 m, 14°48' S, 60°23' W, 4 April 1993, Arroyo et al. 220 ( UC) ; BRAZIL. Amazonian Region , 30 April 1884, Oyster s.n. (P) ; COLOMBIA. Boyacá: Carretera Chiquinquirá a Pauna , 1700 m, 13 October 1967, Mejía et al. 3567 ( NY) ; Santa Marta , 1898, Smith 2715 ( NY) ; Valle: Yotoco, Corregimiento Jiguales , Finca La Camelia , 1450 m, 15 January 1985, Wilson 879 ( UC) ; ECUADOR. Morona-Santiago: Gualaquiza , 800 m, 3°26' S, 78°31' W, 31 July 1993, Fay & Fay 4206 ( NY) ; Zamora-Chinchipe: between San Pablo and Nabija , 1465 m, 04 o 02’18’’S, 78 o 47’52’’W, 23 July 2004, Croat 91981 ( UC) ; PERU. Cajamarca: San Ignacio, Camacá , 1250–1800 m, 5°04' S, 78°55' W, 7 March 1997, Campos & Corrales 3509 ( NY) ; San Martin: Lamas , District Lamas, 01 October 1937, Belshaw 3503 ( NY) ; VENEZUELA. Barinas: Bolívar, 900 m, 8°48' N, 70°32' W, 19 November 1982, Smith et al. 1367 ( UC) ; Mérida, 2000 m, 1893, Mocquerys s.n. (F) ; Falcón: Sierra de San Luis, Montaña de Paraguariba , 1400 m, 23 May 1979 , Falcón 668 ( UC) ; Lara: Palavecino , 500 m del sector urbanizado, 1300 m, 09 o 55’N, 69 o 17’W, 28 September 1984, Rivero 720 ( UC) ; Yaracuy: Limites Distrito Nirgua-Distrito San Felipe , 1000–1200 m, 10°14'00" N, 68°37'30", 24 March 2005, Meier et al. 11342 ( UC) .

Habitat and distribution:—Terrestrial in mountain rainforest, 800–1500 m. Mesoamerica ( Mexico, Honduras, Nicaragua and Costa Rica), West Indies (Hispaniola) and South America ( Venezuela, Colombia, Ecuador, Peru, Brazil and Bolivia; Fig. 23C ; Tab. 01).

Notes:— Ctenitis microchlaena is similar and commonly confused with C. submarginalis , differing from it by segment apex rounded, basal veins from adjacent segments end at margin well above the sinus, indusia always present and conspicuous, costa and abaxial laminar surface with dense catenate trichomes ( Fig. 21C, E, F ). While C. submarginalis segment apex is apiculate ( Fig. 26A ), the basal veins from adjacent segments usually end at margin or somewhat above the sinus, but sometimes well above the sinus, the indusia can be absent, small and inconspicuous or large and conspicuous (plants from Argentina, Paraguay, Uruguay and southern Brazil), with or without sparse catenate trichomes on costa and abaxial laminar surface. Generally, C. microchlaena is more pilose

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than C. submarginalis , with catenate trichomes, which density is similar to C. falciculata . However, C. falciculata rachis scales are dark brown to blackish and to 2.1 mm long, while C. microchlaena rachis scales are castaneous and 2.4–5.0 mm long, and these two species are not sympatric.

Ctenitis microchlaena is firstly recorded to Brazil, based on two old collections. One in P (P01415555!) labeled as from Brazil, near to Amazonian river. Such sample is small, seems to be from a young plant and has the basal pinnae reduced, a feature that may be due to early development, also observed in some small specimens of C. microchlaena from Peru, Colombia and West Indies. The other collection is in F (Luerssen s.n. —F 2002990!), it is from a plant that was cultivated at Leipzig Botanical Garden, which origin reported on the label is Brazil, without further information.

Christensen (1913a) considered this taxon as Dryopteris karstenii , supposing that A. microchlaena (which type he has not seen) should be D. submarginalis ( C. submarginalis ). Later, dealing with pteridophytes from Hispaniola Christensen (1936) suggested that D. karstenii was a weakly marked species, perhaps better to be united with D. submarginalis . As already mentioned, these taxa are similar, but the differences pointed by some authors ( Christensen 1913 a, Tryon & Stolze 1991, Moran 1995, Mickel & Smith 2004) and observed by us are quite constant.

Fée (1857) described Aspidium microchlaena based on the collection Schaffner 459 from Mexico. As we known, Fée did not designate a holotype neither specified a herbarium. Tryon & Stolze (1991) and Mickel & Smith (2004) presumed that the holotype was in P, but they informed that this collection was not found there, and these authors indicated an isotype in K. In fact, there is no Schaffner 459 in P. Fournier, a French botanist, which worked in P, cited this collection as A. microcarpon ( Fournier 1872) . Probably, Fournier may have been the last to examine this material, or he cited it without examining. However, the only sheet of the collection cited by Fée (1857) is in K and this one may be the lectotype. As Tryon & Stolze (1991) were the first who reported that a supposed holotype was not found in P and cited an isotype in K, the lecotypification is attributed to them, although inadvertent (Art. 7.10, 9.9 and 9.19 of ICN — McNeill et al. 2012).

Braun (1858) described Aspidium karstenii based on a cultivated plant in Horto Berolinense, from spores of other plant collected in Venezuela by Karsten & Moritz. He also mentioned other syntypes: Moritz 209 and Gollmer s.n. Christensen (1913a), Tryon & Stolze (1991) and Mickel & Smith (2004) cited that the type (or holotype) was Karsten s.n. in B, probably because most specimens studied by Braun are in B. The only sheet in B with a label handwritten by Braun as A. karstenii is Gollmer s.n. Considering all this, we decided to designate this Gollmer s.n. as the lectotype of A. karstenii (Art. 9.2, 9.5, 9.11, 9.12 of ICN — McNeill et al. 2012).

Fée described Aspidium obtusilobum and cited the collection Schaffner 213, at the same page of A. microcarpon (see “Names of Uncertain Application”), three pages after describing A. microchlaena ( Fée 1857) . Christensen did not see such material and treated A. obtusilobum as an unknown species. In addition he erroneously cited Schaffner 105, which corresponds to the page of its description not to the collecting number ( Christensen 1913a). Smith (1981), Mickel & Beitel (1988) and Mickel & Smith (2004) followed Christensen’s error and presumed that the type should be in P, but it had not been found there. Indeed, in P there is no Schaffner 105 or 213, however there is a sheet of Schaffner s.n. (P00642678!) from Mexico without further information about the locality, on it is written “ Aspidium obtusilobum ”, but there is no Fée’s label. Smith (1981) and Mickel & Smith (2004) have already mentioned that such sheet could be a possible uncited original material. We found a sheet of Schaffner 213 in RB, on its label is written “ Mexique, Huatusco”, exactly as in the protologue. Some ferns specimens described by Fée once belonged to Pedro II, emperor of Brazil. After his death, they became property of M. Cosson in Paris and later were incorporated in P ( Underwood 1905). Some specimens remained in Brazil at Jardim Botânico do Rio de Janeiro, where is the RB herbarium. Then, we designate the sheet in RB for lectotype. Mickel & Smith (2004) and the website tropicos.org, present the name A. obtusilobum Fée as an illegitimate name, due to the earlier homonym A. obtusilobum Willd. , without reffering any page to this supposed Willdenow’s name ( Willdenow 1810). However, in Willdenow (1810) there is no such name. The most similar names in Willdenow (1810) are A. obtusifolium (page 231), A. obtusatum (241) and A. obtusum (254). Then, according to Art. 53.1 and 53.3 of ICN ( McNeill et al. 2012) the name under Fée’s authoring is not illegitimate.

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Figures

FIGURE 21. Some morphological characters of Ctenitis from South America. A–B: Ctenitis laetevirens (Schimitt 561, BHCB). A: abaxial surface of portion of pinna. B: abaxial costa scale. C–F: Ctenitis microchlaena (unknown s.n., UC). C: abaxial surface of a portion of rachis and two basal basiscopic segments of a medial pinna. D: abaxial costa scale. E: abaxial surface of a pinna, showing indument. F: sorus with indusium. Scale bars A, B, D e E = 1 mm; C = 1 cm; F = 0.5 mm

FIGURE 23. Distribution of four species of Ctenitis in South America. A: C. laetevirens. B: C. megalastriformis. C: C. microchlaena. D: C. nervata.

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.