A taxonomic monograph of the fern genus Ctenitis (Dryopteridaceae) in South America Author Viveros, Raquel Stauffer Universidade Federal de Minas Gerais, Instituto de Ciências Biológicas, Departamento de Botânica, Caixa Postal 486, 30123 - 970, Belo Horizonte, Minas Gerais, Brazil Author Rouhan, Germinal Muséum National d’Histoire Naturelle, UMR 7205 CNRS-MNHN-UPMC-EPHE, Institut de Systématique, Evolution, Biodiversité - ISYEB, Sorbonne Universités, Herbier National, 16 rue Buffon, CP 39, F- 75231 Paris cedex, France Email: raquel. s. viveros @ gmail. com raquel.s.viveros@gmail.com Author Salino, Alexandre Universidade Federal de Minas Gerais, Instituto de Ciências Biológicas, Departamento de Botânica, Caixa Postal 486, 30123 - 970, Belo Horizonte, Minas Gerais, Brazil text Phytotaxa 2018 2018-01-23 385 1 448 450 http://dx.doi.org/10.11646/phytotaxa.385.1.1 journal article 10.11646/phytotaxa.385.1.1 1179-3163 13705270 2. Ctenitis ampla (Humb. & Bonpl. ex Willd.) Ching (1940: 250) . Figs. 03C, 06A. 08C–D, 09A, 12C–D, 14B. Polypodium amplum Humboldt & Bonpland ex Willdenow (1810: 207) . Aspidium amplum (Humb. & Bonpl. ex Willd.) Mettenius (1858: 74) , non (Humb. & Bonpl. ex Willd.) Christ in Bommer & Christ (1896: 214) , nom. illeg. Nephrodium amplum (Humb. & Bonpl. ex Willd.) Baker in Hooker & Baker (1868: 285) . Type:— VENEZUELA . Caripe, Humboldt s.n. ( holotype B–W-19722-010!). Polypodium sloanei Poeppig ex Sprengel (1827: 59) . Dryopteris sloanei (Poepp. ex Spreng.) Kuntze (1891: 813) . Ctenitis sloanei (Poepp. ex Spreng.) Morton (1969: 66) . Type: — CUBA . Poeppig s.n. ( lectotype L, designated by Morton 1969, isolectotypes B 20 0052237!, B 20 0052238!, B 20 0052239!, B 20 0052240!, B 20 0052241!, B 20 0052242!, BM 000605200!, BR 0000006989132!, BR 0000006988487! E 00106040!, F 2009038!, HAL 86821 [image!], HBG, K 000590320!, NY 02006034!, P 00642684!, P 00642685!, P 00642686!, P 00642687!, P 00642688! US 00810676!). Aspidium nemophilum Kunze (1834: 95) . Dryopteris nemophila (Kunze) Christensen (1920: 57) . Ctenitis nemophila (Kunze) Ching (1940: 250) . Type: — PERU . Pampayaco. July 1829 ( holotype LZ destroyed); neotype : — PERU . Huanuco : Muña, in dry woods, 23 May–04 June 1923 , Bryan 530 (F 1829440!, designated here ). Aspidium catocarpum Kunze (1834: 95) . Dryopteris catocarpa (Kunze) Kuntze (1891: 812) . Ctenitis catocarpa (Kunze) Morton (1951: 12) . Type: — PERU . inter Cassipi et Papayaco ( holotype LZ, destroyed); neotype : — PERU . San Martin : San Martin , 15 km E of Sharpaja on road to Chazuta, along Quebrada Chumia near Mal Paso Chumia on Rio Huallaga, tropical wet forest and second growth along stream, 76 o 10’W 06 o 36’ S , 04 August 1986 , Knapp 7869 (MO 3698517!, designated here , isoneotype NY!). Aspidium furcatum Klotzsch (1847: 371) . Dryopteris furcata (Klotzsch) Kuntze (1891: 81) . Type: — COLOMBIA . Moritz 37 ( lectotype BM 000937870!, designated here ). Polypodium paleaceum Hooker (1847: 166) , nom. illeg., non Borkhausen (1798: 20) , non Powell ex Baker in Hooker & Baker (1874: 505) . Type: — ECUADOR . Charles and James Islands, Darwin s.n. ( lectotype CGE, designated by Porter 1980 ). Nephrodium palatanganum Hooker (1862: 260) . Dryopteris palatangana (Hooker) Christensen (1920: 56) . Ctenitis palatangana (Hooker) Ching (1940: 250) . Type: — ECUADOR . Palatanga. Spruce 5256 ( holotype K 000200145!). Aspidium culcita Christ (1906: 54) . Dryopteris culcita (Christ) Christensen (1913b: 31) . Type: — COSTA RICA . Wercklé s.n. ( lectotype P 00642689! designated here , isolectotypes BM 000937864!, MO 1638975!, US 00067014!). Stems erect, 3.7–5.7 cm diam., scales 17.9–32.5 × 0.1–0.3 mm , light castaneous or castaneous, clathrate, linear, entire, without fimbriae; leaves 80–200 cm long; petioles 30–83 cm × 4.4–10.6 mm , with 6–10 vascular bundles at base, stramineous or tan, scales 7.21–25.34 × (0.12) 0.18–0.68 (1.54) mm, light castaneous or castaneous, clathrate, tangled at base, becoming ascending towards distal portion, flattish, flaccid, linear with truncate or slightly cordate base and filiform apex, entire, without fimbriae, catenate trichomes absent abaxially, sparse glandular trichomes or absent; laminae 50–117 × 40–120 cm , width wider than 2/3 of length or more, 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 10–14 pairs, the basal ones stalked to 25.5 (51.2) mm long, the medial ones stalked to 15.3 mm long, the apical ones stalked to 7.2 mm long or sessile, basal pinnae basiscopically enlarged, the medial 32.5–44.5 × 10–18 cm , lanceolate, apex attenuate; adaxial pinnae axes with sparse or dense scales on pinna rachis 0.9–3.4 × 0.1–0.4 mm , stramineous or castaneous, lanceolate, catenate trichomes dense on pinna rachis and costa, sparse on costule, rare on veins, bacilliform trichomes absent; adaxial laminar surface between veins glabrous or with sparse catenate trichomes near margins; abaxial pinnae axes with scales sparse or dense on pinna rachis, costa and costule, (0.9) 1.3–4.3 (5.6) × 0.1–0.7 mm , light castaneous or castaneous, clathrate, patent or ascending, flattish or sometimes vaulted at base, flaccid, lanceolate with cordate base and filiform apex, entire or slightly denticulate, with or without some short fimbriae at base and laterally, catenate trichomes sparse on pinna rachis and costa, sometimes on costule and veins, bacilliform trichomes sparse on costa, costule and veins, glandular trichomes sparse or absent on pinna rachis, costa, costule and veins, filiform trichomes absent; abaxial laminar surface between veins glabrous or with sparse catenate and bacilliform trichomes; pinnules 11–19 pairs, 24 • Phytotaxa 335 (1) © 2018 Magnolia Press VIVEROS ET AL. A TAXONOMIC MONOGRAPH OF CTENITIS IN SOUTH AMERICA FIGURE 13. Some morphological characters of Ctenitis from South America. A–E: Ctenitis aspidioides ( Salino 14908, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: sorus with indusium. C: detail of abaxial surface of a medial pinna showing indument. D: petiole scale. E: abaxial costa scale. F–H: C. deflexa ( Salino 2383, BHCB). F: abaxial surface of a portion of rachis and two basal segments of a medial pinna. G: rachis scale. H: abaxial costa scale. Scale bars in A = 1 cm; B = 0.5 mm, C – H = 1 mm. Phytotaxa 335 (1) © 2018 Magnolia Press • 25 the basal ones stalked to 9.1 mm long, the medial ones stalked to 4.3 mm long or sessile, the apical sessile, the medial 6.0–13.3 × 1.31–3.3 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 attenuate or acute; ultimate segments 12–22 pairs, (1.7) 2.5–8.1 mm wide, entire or serrate, apex rounded or acute, margin with catenate trichomes, the distance from each other is narrower than segments width; veins simple or 1–2- forked, 6–9 pairs (before forking) per segment, the basal ones from adjacent segments end at margin well above the sinus; sori medial, indusia absent, or small and incospicuous or large and conspicuous, entire, with bacilliform trichomes; spores with coarse, irregular echinae. Selected specimens examined :— ARGENTINA . Salta : Rivadavia , Cabeceras del Santa María , 18 June 1944 , Willink 215 ( GH ) ; BOLIVIA . Beni : Moxos , Chimanes Forest , 260 m , 15°10' S , 66°37' W , 25 July 1990 , Fay & Fay 2810 ( MO ) ; Yacuma , campamento Campo Monos , bajando por el río Curiraba , 195 m , 14º38’S , 66º04’W , 17 November 1996 , Acebey 42 ( UC ) ; Chuquisaca : Hernando Siles , Área protegida municipal de Monteagudo , 1300 m , 19°48' S , 63°57' W , 8 June 2000 , Huaylla & Wendelberger 7 ( UC ) ; Luis Calvo , Comunidad Chuya Yaco , 1239 m , 19°45'12" S , 63°52'36" W , 20 October 2005 , Lliully et al. 476 ( MO ) ; Tomina , aprox. a 50 min. bajando de Abra Santa Cruz , 1345 m , 19º18’57.3”S , 64º05’15.1”W , 12 October 2004 , Gutiérrez 1011 ( NY ) ; La Paz : Abel Iturralde , Parque Nacional Madidi , campamento de guardaparques Sadiri , caminho Sadiri-San José de Uchupiamonas , 740 m , 14º10’S , 67º55’W , 4 July 2004 , Jimenez 2561 ( UC ) ; Franz Tamayo , Parque Nacional Madidi , 282 m , 14°11'16" S , 68°20'04" W , 11 April 2002 , Paniagua 4226 ( UC ) ; Nor Yungas , Coroico ( Polo-Polo bei Coroico ), 16°11'38" S , 67°43'55" W , October 1912 , Buchtien ( UC ) ; Santa Cruz : Florida , Localidad de Bella Vista , 1359 m , 18°16'06" S , 63°40'24" W Alt : 1359 m , 28 November 2008 , Calzadillaet al. 392 ( MO ) ; Sara , Cerro del Amboró , 1000 m , 17°43'56" S , 63°39'05" W Alt : 1000m , 19 October 1916 , Steinbach 2984 ( GH ) ; Vallegrande , 1200 m , 18º47’S , 63º51’W , 20 May 1996 , Kessler et al. 5926 ( UC ) ; Velasco , Serrania de Huanchaca , 200–300 m , 14°37' S , 60°42' W , 4 November 1991 , Foster 13694 (F) ; BRAZIL . Ceará : Pacatuba , Serra de Pacatuba , ca. 600 m , 03 o 58’S , 38 o 47’W , 22 july 1997 , Almeida – Neto & Meirelles 270 ( HUEFS ) ; Minas Gerais : Januária , Vale do Peruaçu , mata ciliar a caminho do janelão, 15º07’23”S , 44º13’34”W , 20 July 1997 , Salino 3257 ( BHCB ) ; COLOMBIA . Cundinamarca : Gachetá , En el Cemeterio , 1750 m , 4°55'15" N , 73°36'48" W , 18 May 1974 , Acosta-Arteaga 403 ( NY ) ; Risaralda : Pereira, Hacienda Alejandría , 11 February 1990 , Silverstone – Sopkin et al. 5893 ( UC ) ; Santa Marta : Las Vegos Forest , 2500 f, 11°18'26" N , 73°59'11" W , March 1936 , Bennett 27 (F) ; Valle : El Cerrito , Hacienda El Hatico , 1000 m , 3°39' N , 76°19' W , 16 December 1994 , Silverstone-Sopkin & Paz 7162 ( UC ) ; ECUADOR . Andes , 1857, Spruce 5256A ( NY ) ; Cotopaxi : Quevedo-Latacunga road, km 46 from Quevedo , ca. 600 m , 00º55’S , 79º11’W , 1973, Homl-Nielsen 2904 ( UC ) ; Galapagos : Santa Cruz island, 430 m , 3 February 1964 , Itow, S. 95 ( CAS ) ; Morona–Santiago , Morona , Cordillera de Cutucú , 600 m , 02º32’S , 77º54’W , 22 January 2002 , Palacios et al. 15792 ( UC ) ; Pichincha : Hotel Tinalandia , casi 25 km al este desde Sto. Domingo de los Colorados , Cordillera Occidental , 1000m , 8 January 1984 , Moran 3542 ( NY ) ; Tungurahua : Baños , Downhill , 1450 m , 1°24' S , 78°17' W , 8 July 1992 , Fay , & Fay 3568 ( NY ) ; PERU . Huánuco : Leoncio Prado , 1400 m , 6 June 1981 , Young & Sullivan 865 (F) ; Junín : Tarma , Valle del río Chanchamayo , 1500 m , 1918, Esposto 671 ( GH ) ; Madre de Dios : Manu , Parque Nacional del Manu , 350 m , 11°50' S , 71°25' W , 10 August 1984 , Foster 9802 (F) ; Pasco : Oxapampa , Pozuzo , Parque Nacional Yanachanga–Chemillén , 1100 m , 10º10’58’’S , 75º34’25’’W , 23 July 2006 , Monteagudo et al. 12551 ( MO ) ; San Martin : Huallaga , Abajo de La Morada , 1900–2000 m , 6°57' S , 77°32' W , 10 August 1997 , Quipuscoa & Bardales 935 (F) ; VENEZUELA . Anzoátegui : 800–1000 m , 1 March 1945 , Steyermark 61279 (F) ; Aragua : Tovar , 1854, Fendler 204 ( GH ) ; Lara : Torres , Via El Jabón , sítio denominado La Raya , 1500 m , 22 January 1985 , Rivero 86613 ( UC ) ; Margarita Island , 8 February 1901 , Miller & Johnston 169 ( NY ) . Habitat and distribution :—Terrestrial in rainforest in low or highlands, 0–2000 m . United States of America (Florida), Mesoamerica ( Mexico , Honduras , Nicaragua , El Salvador , Costa Rica and Panama ), West Indies ( Cuba , Jamaica , Hispaniola and Trinidad ) and South America ( Brazil , Venezuela , Colombia , Ecuador , Peru , Bolivia and Argentina ; Fig. 14B ; Tab. 01). Notes :— Ctenitis ampla is a 2–4-pinnate-pinnatifid species (Fig. 03C), similar to C. equestris var. equestris , C. grisebachii and other decompound lamina species from Mesoamerica and West Indies. Ctenitis ampla and C. equestris var. equestris scales are flaccid, clathrate, flattish or vaulted with cordate base, with or without some short fimbriae at base and laterally, but the C. ampla ones are uniformly light castaneous or castaneous (Fig. 06A–B), while C. equestris scales are brownish with pale edges ( Fig. 17B ), often iridescent. Ctenitis grisebachii has stiff uniformly castaneous or brown, always flattish scales with truncate base and without fimbriae ( Fig. 19G ), the ones along petiole and rachis are usually retrorse. 26 • Phytotaxa 335 (1) © 2018 Magnolia Press VIVEROS ET AL. A TAXONOMIC MONOGRAPH OF CTENITIS IN SOUTH AMERICA FIGURE 14. Distribution of four species of Ctenitis in South America. A: C. abyssi . B: C. ampla . C: C. anniesii . D: C. aspidioides . Phytotaxa 335 (1) © 2018 Magnolia Press • 27 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. Ctenitis ampla is morphologically variable and widely distributed. Because of that, some authors considered it as one of the species complex in Ctenitis , justifying the need of a revision and highlighting the importance of examination of all types and greater amount of materials from all its distribution range ( Christensen 1920 , Tryon & Stolze 1991 ). Christensen (1920) , despite manifesting be in doubt, considered C. sloanei ( Dryopteris sloanei ) under C. ampla ( D. ampla ) and distinguished the Andean form as D. nemophila . Later, Morton (1969) pointed that C. ampla should be what Christensen (1920) referred as D. nemophila , distinguishing it from D. sloanei and proposing the new combination of it under Ctenitis . His major arguments were based on the holotype photograph of Polypodium amplum and on the collection Fendler 204 from Venezuela , which Christensen (1920) cited as D. nemophila . Since then, the specimens from United States , Mexico , Central America and West Indies were usually called C. sloanei and most specimens from South America were called C. ampla . The only and most recent taxonomic treatment that dealt with these supposed two species was Tryon & Stolze (1991) for flora of Peru . These authors also were uncertain about the application of both names or even if more taxa should be considered among this complex. Indeed, the Andean specimens, especially on high elevations from Peru and Bolivia , are the most similar to the holotype of P. amplum than to P. sloanei . They are quite larger and robust plants, the scales and the laminae are darker (castaneous), and the segments are wider and almost glabrous (less catenate trichomes at margin, less bacilliform trichomes on veins abaxially). Even with the reasoning attempts of Morton (1969) and Tryon & Stolze (1991) to distinguish C. ampla from C. sloanei , we consider only one variable species, under the name C. ampla . After analyzing numerous exsiccatae from all known distribution range, we realized that several characters overlap, which render infeasible to distinguish two taxa. Such characters as the color of bacilliform 28 • Phytotaxa 335 (1) © 2018 Magnolia Press VIVEROS ET AL. trichomes (reddish or yellowish), if they are few or many, the color of scales (castaneous or light castaneous) or how conspicuous the indusia are, or how wide are the segments. Furthermore, on specimens with or without (or inconspicuous) indusia from different regions, in only one sorus there are sporangia capsule glabrous or with one or more bacilliform trichomes (Figs. 08C–D), a feature seen only in this species among all the others Ctenitis from the New World. The spore surfaces of different “morphotypes” are also the same. Maybe the quite different morphology is result of different ploidy, what was not investigated in this work, but is certainly an interesting issue to explore. Another name recognized by Christensen (1920) that we consider as a synonym of C. ampla is Dryopteris palatangana . Christensen (1920) was based only on the original material of Nephrodium palatanganum . We have examined some specimens from Ecuador , inclusive from Palatanga, and they are just less scaly plants, which scales are castaneous. The specimens from Galapagos Islands are the smallest plants, consequently the segments are smaller and usually acute at apex, the scales are lighter and dense on axes, and there are dense yellowish bacilliform trichomes along throughout the veins abaxially. Some old collections with these features are identified as Polypodium paleaceum . The ones from Costa Rica and Panamá , as Aspidium culcita , are quite the same, but the laminae are darker and folded, probably due to preparation process. In the original description of Polypodium sloanei , no type neither herbarium was indicated ( Sprengel 1827 ). Later Kunze (1834) mentioned a material that was in “Herb. Poep. Kze”, which is currently LZ herbarium. After this, other authors cited the holotype of P. sloanei as destroyed in LZ and some isotypes in other herbaria ( Tryon & Stolze 1991 , Mickel & Smith 2004 ). Indeed, the material in LZ was destroyed during the Second World War. Morton (1969), however, did not mention a holotype , but an isotype in L. What Kunze did must be considered as an inadvertent lectotypification (Art. 9.9. of ICN— McNeill et al. 2012 ). Nonetheless, as the specimen in LZ was destroyed, the isotype cited by Morton (1969) must be considered as lectotype , and its several duplicates as isolectotypes . Unfortunately, Polypodium nemophilum and P. catocarpum ’s type materials were also in LZ, thus also destroyed. No other author resolved the typification. No specimens that could be original material of these names were found. Then we designate neotypes for them (Art. 9.16 of ICN— McNeill et al. 2012 ), that represent Christensen’s morphology concept of Dryopteris nemophila ( Christensen 1920 ) , in our understanding, and they are from Peru , as the ones in original description. We also propose the lectotypification of Aspidium furcatum , once no original material was found in B. Probably it must have been destroyed as other types of names described by Klotzsch. The sheet chosen for lectotype (Art. 9.12 of ICN — McNeill et al . 2012 ) is a fragment of the holotype . Such fragment is in BM, purchased from Christensen’s herbarium. For Aspidium culcita , as we have examined four sheets in different herbaria of the collection mentioned in its protologue ( Wercklé s.n. ), we designate as lectotype (Art. 9.12 of ICN — McNeill et al . 2012 ) the sheet in P, which is from Christ’s herbarium, the author of this name.