Ctenitis equestris (Kunze) Ching (1940: 250) var. equestris

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

9. Ctenitis equestris (Kunze) Ching (1940: 250) var. equestris . Figs. 09I, 17A–B, 18B. Aspidium equestre Kunze (1844: 347) . Dryopteris equestris (Kunze) Christensen (1920: 54) . Type:— MEXICO. Leibold 42 (holotype LZ destroyed, lectotype B 20 0055411!, designated here, isolectotype HBG).

Polypodium alsophiloides Liebmann (1849: 208) . Type:— MEXICO. Oaxaca: “Trapiche de la Concepcion”, Liebmann sn. [Pl. Mex. 2406] (lectotype C designated by Smith 1981, isolectotype B 20 0055237!).

Lastrea ciliata Liebmann (1849: 273) . Type:— MEXICO. Veracruz: Near S. Antonio Huatusco; 1450 ft, Liebmann s.n. [Pl. Mex. 2394, Fl. Mex. 199] (C—two sheets; US 00067013!) .

Aspidium bourgeaui Fournier (1872: 98) (“ bourgaei ”). Dryopteris bourgeaui (E. Fourn.) Christensen (1906: 255) . Type:— MEXICO. Veracruz: Vallée de Cordova, Bourgeau 1839 (lectotype P00642730!, designated here, isolectotypes BM 000937868!, BR 000000698770!, K 000590296!, K 000590297!, LE!, MPU 15242 [image!], MPU 15243 [image!], P 00642729!, P 00642731!, US 00067006!, US 00067007!, MICH 1190030 [image!]).

Aspidium scabriusculum Davenport (1896: 255) , nom. illeg., non Mettenius in Salomon (1883: 69). Dryopteris davenportii Christensen (1906: 260) . Type:— MEXICO. Veracruz: Ravines above Orizaba, 6 February 1895, Pringle 6132 (lectotype GH 00020486!, designated here, isolectotypes B 20 0055234!, BM 000788116!, CAS 0000779!, CM 0113 [image!],

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COLO 00348128 [image!], K 000590295!, F 0075886F!, F 0075887F!, GH 00967126!, GH 00967127!, GOET 008719 [image!], MEXU 01223467 [image!], MEXU 01223466 [image!], MEXU 01223465 [image!], MO 102952!, NDG 04227 [image!], NY 00149420!, NY 00149427!, P 00642725!, P 00642726!, P 00642727!, P 00642728!, PH 00004818 [image!], PR 4585!, S-P-8918!, UC 123306!, US 00067042!, VT UVMVT015729 [image!], WU 032301 [image!]).

Stems erect or ascending, 2.0–4.0 cm diam., scales 8.9–22.6 × 0.3–1.1 mm, light castaneous or castaneous, clathrate or subclathrate, linear, entire, without fimbriae; leaves 30–150 cm long; petioles 30–75 cm × 2.6–8.0 mm, with 5 vascular bundles at base, stramineous or tan, scales 5.55–15.81 × 0.43–1.50 mm, light castaneous or castaneous, clathrate, tangled at base, becoming ascending and imbricate towards distal portion, flattish, flaccid, linear with truncate base and filiform apex, entire, with or without some short fimbriae at base, sparse catenate trichomes abaxially, sparse glandular trichomes; laminae 25–80 × 30–90 cm, width wider than 1/2 of length, 3–4- pinnate-pinnatifid basally, 2–3-pinnate-pinnatifid medially and apically, deltate, apex confluent; rachises more or less straight, stramineous or tan, scales like those on distal portion of petioles, sparse catenate trichomes abaxially, sparse glandular trichomes; pinnae 9–15 pairs, the basal ones stalked to 25.8 mm long, the medial ones stalked to 24.0 mm long, the apical ones stalked to 2.1 mm long, basal pinnae basiscopically enlarged, the medial (10.5) 17–50 × (4) 8.5–40 cm, lanceolate, apex attenuate or caudate; adaxial pinnae axes with sparse scales on pinna rachis 0.5–2.0 × 0.1–0.4 mm, brownish, somewhat iridescent, black-walled in middle with pale edges, lanceolate or ovate, catenate trichomes dense on pinna rachis and costa, sparse on costule and veins, bacilliform trichomes absent; adaxial laminar surface between veins glabrous or with sparse catenate trichomes; abaxial pinnae axes with scales sparse on pinna rachis and costa, rare on costule, 0.8–3.1 × 0.2–0.6 mm, dark brown to blackish, sometimes pale at edges, iridescent, clathrate, appressed or imbricate, flattish or sometimes vaulted at base, flaccid, lanceolate with cordate base and filiform or attenuate apex, entire or slightly denticulate, with or without some short fimbriae at base, catenate trichomes sparse or dense on costa, sparse or rare on costule and veins, bacilliform trichomes absent or sparse on costa, costule and veins, glandular trichomes sparse on pinna rachis, costa and costule, rare on veins, filiform trichomes absent; abaxial laminar surface between veins glabrous or with sparse catenate and bacilliform trichomes near margins; pinnules 7–13 pairs, the basal ones stalked to 5.3 mm long, the medial ones stalked to 0.7–1.8 mm long, the apical sessile, the medial (2.3) 3.8–10.5 × (0.5) 1.1–3.3 cm, lanceolate, incised more than 3/4 of the distance between the segment apex and costa, almost pinnatisect, pinnatisect basally, basal segments as long or longer than the next, apex acute, attenuate or caudate; ultimate segments 8–14 pairs, 1.6–7.5 mm wide, entire, crenate or serrate, apex rounded or acute, margin with catenate trichomes, the distance from each other is narrower than segments width; veins simple or 1–3-forked, 6–9 pairs (before forking) per segment, the basal ones from adjacent segments end at margin well above the sinus; sori medial, conspicous, entire, with bacilliform trichomes; spores with coarse, irregular echinae.

Selected specimens examined:— COLOMBIA. Santander: western slope of Mount Peña Blanca, near Charta , 2500–2600 m, 11 February 1927, Killip & Smith 19276 ( GH) ; ECUADOR. Galapagos: Albemarle Island , 27 August 1905, Stewart 959 ( NY) ; VENEZUELA. Distrito Federal: Cerro El Avila , Parque Nacional El Avilla , 1990 m, 10°32' N, 66°52' W 10 February 1992, Meier 1641 ( UC) ; Lara: Trujillo, Carache, via Cerro Gordo , 2100 m, 09º32’N, 70º09’W, 21 May 1987, Rivero 1178 ( UC) .

Habitat and distribution:—Terrestrial in montane rainforests, 500–2600 m. Mesoamerica ( Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama) and South America ( Venezuela, Colombia and Ecuador — Galapagos; Fig. 18B ; Tab. 01).

Notes: —Among the other 2–4-pinnate-pinnatifid (Fig. 03C) South American species, C. equestris var. equestris differs by its scales dark brown to blackish sometimes pale at edges and iridescent ( Figs. 17A – B ). This taxon is recorded for the fist time in South America, through some herbaria specimens previously identified as C. ampla . Nonetheless, this taxon is widely distributed in Mesoamerica. The other variety under this species is C. equestris var. erosa ( Stolze 1977: 41) , which occurs in Mexico, Guatemala and Honduras.

The holotype indicated in the protologue of Aspidium equestre was in Römer’s Herbarium, which was incorporated by LZ, and then destroyed during the Second World War. A duplicate with sample of stem and a complete lamina is in B (from Brehmer’s Herbarium) and we designate it to be the lectotype (Art. 9.12 of ICN— McNeill et al. 2012). There is another in HBG ( Mickel & Smith 2004), which we have not examined in hands.

Liebmann (1849) was unaware about the type material of A. equestre and described the same taxon as Polypodium alsophiloides and Lastrea ciliata , two morphological extremes, but still the same taxon.

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Aspidium bourgeaui was described by Fournier (1872) as distinct from A. equestre , by subtle differences in segments size and form. As Fournier cited a gathering, Bourgeau 1839, without specifying any herbarium, and there are many duplicates of this collection in several herbaria, we selected one of the sheets in P, where Fournier worked, to be the lectotype (Art. 8.3, 9.2, 9.5, 9.11, 9.12 of ICN— McNeill et al. 2012). The sheet chosen contains a larger fragment of leaf than the others.

In the protologue of Aspidium scabriusculum, Davenport (1896) cited the collection Pringle 6132. Pringle’s collections are widely spread in several herbaria. We found 26 herbarium sheets of Pringle 6132. Mickel & Smith (2004) cited the one in GH as the holotype and other herbaria for isotypes. Davenport worked in Harvard, however he did not indicate a holotype, nor specified a herbarium ( Davenport 1896). For this reason, we designate here as lectotype the same sheet indicated as holotype by Mickel & Smith (2004), certainly examined by Davenport (see Art. 7.10, 9.2, 9 Note 6, 9.9, 9.11, 9.12, 9.19 and 40 Note 1 of ICN — McNeill et al. 2012).

Figures

FIGURE 17. Some morphological characters of Ctenitis from South America. A–B: Ctenitis equestris (Breedlove 33659, CAS). A: abaxial surface of portion of pinnules and pinna rachis (scale bar = 1 cm). B: abaxial costa scale. C–G: C. eriocaulis (Pietrobom 5007, BHCB). C: abaxial surface of a portion of rachis and two basal segments of a medial pinna. D: adaxial surface of a portion of a segment. E: abaxial surface of a portion of a pinna. F: sorus with indusium. G: abaxial costa scale. Scale bars in A and C = 1cm; B, D, E, G = 1mm; F = 0.5 mm.

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.