Cyathea lasiosora (Kuhn) Domin (1929a: 262)

Lehnert, Marcus, 2016, A synopsis of the exindusiate species of Cyathea (Cyatheaceae-Polypodiopsida) with bipinnate-pinnatifid or more complex fronds, with a revision of the C. lasiosora complex, Phytotaxa 243 (1), pp. 1-53 : 14-19

20. Cyathea lasiosora (Kuhn) Domin (1929a: 262) . ( Figs. 2C , 3 ). Alsophila lasiosora Mett. ex Kuhn (1869: 157) . Alsophila tarapotensis Rosenstock (1909: 291) , nom. superfl. Trichipteris lasiosora (Kuhn) Tryon (1970: 45) . Type:— PERU. San Martin: “Mt. Campana, East Peru,” R. Spruce 4349 [correctly 4249] (lectotype B-20-0000191!, here designated, isolectotypes BM-000937696–000937699!, COL [fragment of GH], GH-00020417!/-00020418!, K-000589978!, LE-00008108!, NY-148693!, P-00631726!/-00631727!), US-00066259! [fragment of K-000589978]!/- 00066288 [fragment ex Herb. Bonaparte]). Exluded because putatively a different sample of the same species [most labelled “ Spruce 4249 var. videtur ”]: B-20-0000190!, K-000589975–000589977!, P-00631728!/-00631729!.

Alsophila nigra Martius (1834: 47) . Trichipteris nigra (Mart.) Tryon (1970: 46) , not Cyathea nigra Linden ex Fournier (1876: 73) . Type:— BRAZIL. Amazonas: “Flumen Japurá in Provincia Rio Negro dicta”, C.F.P. Martius s.n. (holotype M!, isotypes B-20- 0000188!, NY-148719! [fragment of Herb. Martius]!/-148720! [fragment of B], LE-00008117!, M! [3 sheets]).

Alsophila mapirensis Rosenstock (1928: 57) . Type:— BOLIVIA. La Paz: [Larecaja,] San Carlos, region of Mapiri, [ca. 15°24’S, 68°11’W,] 850 m, 9 December 1926, O. Buchtien 292 (lectotype S-R-207!, here designated, isolectotypes NY-00148699!, UC-478278!).

Alsophila killipii Maxon (1942: 58) . Type:— PERU. Loreto: Between Yurimaguas and Balsapuerto, 1929, E.P. Killip & J.D. Smith 28133 (holotype US-00066255!/-00066256!/-00066257!, isotypes F n.v., NY-00148692!).

Trunks to 3 m tall, slender, 2–4(–5) cm diameter, without old petiole bases, with dark gray-brown frond scars 3.0–3.5 × 2.0– 2.5 cm; epidermis densely covered with dark brown, lanceolate scales similar to petiole scales; apices hidden between petiole bases; without adventitious buds. Fronds to 250 cm long, spirally arranged along trunk, not clustered at apices, arching to pendent. Petioles 25–45 cm long, proximally muricate with spines to 1 mm long, dark brown to dark auburn, matte to weakly shiny, with dense scurf consisting of appressed reddish brown trichomidia and dentate or cilitae, ovate to isiodiametic squamules, uniseriate hairs absent; petioles basally without lenticels, or not detectible in dried material; only basally deciduously scaly. Petiole scales lanceolate to ovate lanceolate, to 15.0 × (1.5–)2.0–3.0 mm, thick-textured, bases weakly cordate, basifix to pseudopeltately attached, straight to falcate, apices long acute, flat; scales concordantly bicolorous, dark castaneous to almost black with whitish to brown margins; differentiated margins fragile, the cell rows strongly exserted, with short teeth. Laminae to ca. 100 × 90 cm, bipinnate-pinnatifid, firm herbaceous to chartaceous, matte, dark-green adaxially, often blackish when dried, dark olive green abaxially; apices gradually reduced or abruptly reduced to a rather broad, non-conform section. Rhachises sparsely muricate in proximal parts, otherwise inermous, brown to dark auburn abaxially and adaxially; pubescent with whitish to tan multicellular hairs to 1.0– 1.5 mm long, apressed and persistent adaxially, spreading and thin, ephemeral abaxially, leaving the cortex smooth if abraded; also with reddish brown, scurf similar to petiole scurf. Pinnae to 45 cm long, subsessile or stalked to 0.5–1.0 cm, (7–)12–25 pairs per frond, patent to ascending, alternate, inarticulate, distally narrowly green-alate, distal segments weakly decurrently adnate before ending in a pinnatifid apical section; basal pinna pairs not much smaller than the medial pinnae, weakly reflexed. Costae to 1.5(–2.5) mm wide, inermous, dark ochre to castaneous abaxially, darker adaxially; with whitish to tan, uniseriate hairs 1.0– 1.5 mm long, adaxially appressed, abaxially uniseriate hairs spreading or glabrous, also with scurf of appressed reddish brown trichomidia and branched hairs; junctures of costae and rhachises not or only weakly swollen abaxially, each with an inconspicuous, planar to weakly protruding, elliptic aerophore to 2 × 1 mm, often black when dried. Largest pinnules to 75(–80) × 8–17 mm, lanceolate to linear–oblong, sessile to subsessile, rarely stalked 1–3 mm, inarticulate, 1.0– 1.7 cm between the stalks/costules, truncate to weakly cordate basally, long-acuminate to attenuate apically with serrulate to crenulate margins; costules blackish brown, rarely dark carnose to atropurpureous adaxially and abaxially, adaxially strongly prominent, ridged, and densely hairy with whitish to tan, multicellular hairs, smaller ones antrorsely curved to appressed, to 1.0 mm long, larger ones spreading, to 2.0 mm long, costules abaxially weakly to strongly prominent, densely and persistently hairy with many erect, whitish to reddish brown hairs to 1.5(–2.0) mm long, also with few reddish brown, flat to subbullate squamules to 2.0 mm long; costules basally without pneumathodes, blackened in dried specimens. Largest segments (5.0–)6.5–9.0 × (2.0–)2.5–3.0 mm, segments sessile, adnate, proximal ones patent, distal ones ascending, distally straight or weakly falcate, tips obtuse to rounded, proximal segments usually opposite or nearly so, not shorter than following segments, never remote; sinuses acute to obtuse, to 1.0(–2.0) mm wide; margins mostly crenulated, sometimes subentire or crenate; margins not differently incised in proximal segments of a pinnule; veins planar to weakly protruding adaxially and abaxailly, blackish to dark carnose, ending in the margins; veins adaxially and abaxially with few to many erect, whitish to reddish brown, multicellular hairs to 2.0 mm long on a them, not between the veins; midveins with few, reddish brown bullate squamules to 2.0 × 0.5 mm; sterile veins simple or forked, fertile veins forked. Sori (0.6–)0.8–1.0 mm diameter, ± medial, at vein forks, mature dark orange-brown; indusia lacking; receptacles globose, rather small, 0.2–0.3 mm diameter, paraphyses numerous, whitish to tan, longer than the sporangia (0.5–0.6 mm long), spreading in fresh material but contorted and plastered over the sori when dried, persisting in over-mature sori. Spores white, exospore smooth, finely porate, perispore finely baculate ( Gastony 1979).

Distribution and habitat: — Colombia, Venezuela, Ecuador, Peru, Amazonian Brazil, and Bolivia, in terra firme forest and Andean foothills at 100–1250 m, most common below 1000 m.

Additional specimens examined: — COLOMBIA. Amazonas: Quinche, margen derecha del Rio Caqueta (sur), en frente al Quebrada de Taamanco, [ca. 0°30’51”S, 73°32’28”W,] May 1989, E. Ligia et al. 523 (NY!). Caqueta: Florencia, Cerro La Sardina, [ca. 01°37’N, 75°37’W,] 500 m, 30 March 1940, J. Cuatrecasas 8910 (F). Cauca: Piamonte, Bota Caucana, 01º08.14’39”N, 16º16.55’48”W, 27 July 2002, C. Gonzáles 381 (COL). Vaupés: Río Vaupés, Mitú and vicinity, at base of Cerro Mitú, 27 Sep–20 October 1966, R.E. Schultes et al. 24354 (GH); Vaupés, en Monfort, 30 November 1956, R. Romero Castañeda 3820 (MO!); Mitú and vicinity, lower Río Kubiyú, J.L. Zarucchi & E.W. Davis 1188 (K).— VENEZUELA. Táchira: Cerro Las Minas 18–20 km SE of Santa Ana, 07°36’N, 72°13’W, 1150–1250 m, 11 November 1983, J.A. Steyermark et al. 19848 (UC!).— ECUADOR. Morona-Santiago: Tiwintza, region of Cordillera de Cutucú, N of Río Santiago, Centro Shuar Yapapas, 03º00’31”S, 78º04’09”W, 380 m, 28 October 2005, C. Morales et al. 1440 (MO!, UC!); road Patuca-Santiago, km 44, 03°01’S, 78°12’W, 780–930 m, 12–14 March 1998, B. Øllgaard & H. Navarrete 2879 (AAU!, QCA). Napo: Parque Nacional Yasuní, km 68 de la carretera, 00°43’S, 76°30’W, 270 m, 31 January 2000, H. Navarrete 1520 (AAU!, QCA), parcela de 50 ha, 00°40’S, 76°23’W, 250– 280 m, 31 January 2000, H. Navarrete 1544 (AAU!). Zamora-Chinchipe: Nangaritza Cantón, Pachicutza, trail to Hito, Cordillera del Condor, 04º07’S, 78º37’W, 1200–1300 m, 20 October 1991, W. Palacios 8421 (MO!, UC!), 900 m, 03 December 1990, W. Palacios & D.N. Neill 6502 (MO!); Campamento Miazi, along Río Nangaritza, 900 m, 17 February 1994, H. van der Werff et al. 13167 (MO!, UC!); entrance of the Quimi Valley, more or less 1 km from mining camp, 03°31’41”S, 78°25’33”W, 950 m, 02 November 2004, H. van der Werff et al. 19178 (GOET!, MO!, NY!). Tungurahua: Baños-(Jivaria de) Pintuo (Pastaza-Thal), O. Stübel 995 (B!, NY!).— PERU. Amazonas: Bagua, Distrito Imaza, Yamayakat, 04º55’S, 78º19’W, 350 m, 16 October 1995, E. Rodriguez-R. 514 (MO!, UC!), Comunidad Aguarana de Putuím, 04º55’S, 78º19’W, 680 m, 12 June 1996, E. Rodriguez-R. et al. 977 (MO!, UC!), Condorcanqui, native community Yamayakat, 05º03’24”S, 78º20’17”W, 350 m, March 2006, A. Bonino 403 (MO!), Distr. El Cenepa, comunidad de Mamayaque, Cerro Saake-gaig, 04º34’49”S, 78º14’04”W, 600–800 m, 14 February 1997, E. Rodriguez-R. 1515 (MO!, UC!). Loreto: Loreto, Sierra del Pongo, 800 m, 15 December 1931, Y. Mexia 6270 (MO!); about 2 km N of the village San Antonio at lower Río Marañon, 04º32’S, 73º38’W, 100–200 m, 14 January 1996, H. Tuomisto et al. 8055 (UC!); Río Tigre, just NE of the river, 03º24’S, 74º49’W, 100–200 m, 12 January 2005, H. Tuomisto et al. 14104 (TUR), 2 km S of the village Paicheplaya E of the river, 03º03’S, 75º18’W, 100–200 m, 24 January 2005, H. Tuomisto et al. 14545 (GOET!, TUR), 5 km N of the village Paicheplaya E of the river, 02º59’S, 75º18’W, 100–200 m, 31 January 2005, H. Tuomisto et al. 14763 (GOET!, TUR); Maynas, Panguana primera zona, 3 km E from village, 03°56´S, 73°09´W, 200 m, 30 August 2006, M.J.M. Christenhusz et al. 2154 (GOET), Mishana, Río Nanay, 130 m, 03 December 1977, A. Gentry et al. 21109 (UC!), close to the village of Trece de Fevrero, Km 25 of the road Iquitos-Nauta, 04º04´S, 73º28´W, 100–200 m, 29 December 1994, H. Tuomisto et al. 6614 (GOET!, TUR); Exploration Napo Camp at Río Sucusari, 03º20´S, 72º55´W, 120 m, 19 March 1996, H. van der Werff & Vásquez (MO!, UC!), 20 March 1996, H. van der Werff & Vásquez 13944 (MO!, UC!), 08 April 1997, Vásquez et al. 22762 (MO!, UC!); Maynas, Distr. Sargento Lores, Constancia Norte, 04º07’19”S, 72º55’25”W, 116 m, 11 April 1997, R. Vásquez et al. 22887, MO!, UC!), Distr. Iquitos, Allpahuayo-IIAP, 03º57’19”S, 73º25’47”W, 140 m, 25 April 1997, R. Vásquez et al. 23535 & 23536 (UC!, MO!), Requena, 140 m, 11 August 1988, H. van der Werff et al. 10096, (UC!, MO!). Madre de Dios: Manu, Cerro de Pantiacolla: Rio Palotoa 10–15 km NNW of Shintuya, transect on ridgetop, 12º35’S, 71º18’W, 650–700 m, 12 December 1985, R.B. Foster et al. 10703 & 10722 (F). Pasco: Oxapampa, [as “Prov. Junin ”] Puerto Bermudez, 375 m, 14–17 July 1929, E.P. Killip & J.D. Smith 26550 (NY!); Dist. Palcazú, San Pedro de Pichanaz-Azulis, 10º25’58”S, 75º06’11”W, 670 m, 25 February 2004, L.F. Mellado-N. 720, 734 (MO!); San Francisco de Pichanaz, 10º29’16”S, 75º03’58”W, 770 m, 28 February 2004, L.F. Mellado-N. et al. 927, 928, 929, 930, 931, 1002 (MO!), San Pedro de Pichanaz-San Matías, 10º25’58”S, 75º00’21”W, 1030 m, 1 March 2004, L.F. Mellado-N. & A. Monteagudo 1083, 1085 (MO!), Reserva Comunal Yanesha, Comunidad Nativa Lomalinda-Laguna, sector Azulis, 10º29’32”S, 75º06’30”W, 540–560 m, 20 September 2005, A. Monteagudo et al. 10019 (MO!), sector Nueva Aldea, 10º23’43”S, 75º05’12”W, 620–680 m, 14 October 2005, A. Monteagudo et al. 10605 (MO!), 10º23’13”S, 75º05’28”W, 600–620 m, 17 October 2005, A. Monteagudo et al. 10728 (MO!); Puerto Laguna, 10º18’S, 75º10’W, 400–450 m, 13 September 1984, D.N. Smith 8434 (MO!, UC!). Puno: Río Tavara base camp, 13º21’S, 69º40’W, 400 m, 17 May 1992, A. Gentry et al. 76692 (MO!, UC!). San Martin: Mariscal Caceres, Distrito Tocache Nueve. Puent Palo Blanco Rio Tocache), 10 km west of Tocache Nuevo on road to Puerto Pizana, 08º14’S, 76º35’W, 550–650 m, 12 December 1981, T. Plowman et al. 11354 (GH!, NY!); Uruhuasha, N from Tarapoto (La Banda de Shilcayo), 06°27’S, 76°20’W, 600 m, 3 August 2006, M.J.M. Christenhusz et al. 1966, 1971 (GOET); San Martin, Panguana primera zona, 3 km E from village, 03°56’S, 73°09’W, 200 m, 29 August 2006, M.J.M. Christenhusz et al. 2141 (TUR); San Martin near Tarapoto, 1050–1100 m, 06°26.832’S, 76°17.741’W, 3 Novembr 2010, M. Lehnert 1999 (USM); “prope Tarapoto,” 1855–1856, R. Spruce 4249 (putative type material BM!,K!, TCD!).— BRAZIL. Acre: Cruzeiro do Sul, Rio Jurua & Rio Moa, vicinity of Serra da Moa, 23 April 1975, G.T. Prance et al. 12250 (AAU!, B!); Projecto do Coloniczao Santa Luzia, 45 km from Cruzeiro do Sul to road Cruzeiro do Sul-Rio Branco (BR 364), 07º45’S, 72º15’W, 10–14 September 1985, J. Jangoux et al. 85-004 (UC!). Amapa: Riviere Haut Jari, 02º28’S, 54º46’W, 380 m, 25 August 1993, J.J. de Granville et al. 1246 1 ( US!). Amazonas: Mun. Maráa, Rio Japurá, afluente do Rio Salimoes, ao longo do rio, lugar Maguarizinho, [ca. 01°50’S, 65°22’W,] 29 October 1982, C.A. Cid & J. Lima 3413 (GH!, MO); near mouth of Rio Embira (tributary of Rio Tarauaca), 07º30’S, 70º15’W, 19 June 1933, B.A. Krukoff 4937 (BM!,UC); Río Purus, Rio Ituxi, headwaters of Río Curuqueté, near Ihgarapé João Bento, 100 m, 30 July 1971, Prance et al. 14665 (UC!). Rio Negro: Barra do Rio Negro, R. Spruce 614 (K). Roraima: foothills of Serra Parima, S of Auaris, 04º03’N, 64º22’W, 900 m, 8 February 1969, G.T. Prance et al. 9718 (UC!); vicinity of Posto Surucucu Mission, 02º42–47’N, 63º33–36’W, 19 February 1969, G.T. Prance et al. 10097 (UC!).— BOLIVIA. Cochabamba: Carrasco, Parque Nacional Carrasco, al S del campamento petrolero Ichoa, 17°23’S, 64°24’W, 650m, 21 September 2001, A. Acebey 711 & 718 (LPB!, UC!), Campamento Guacharos, pasando el Río San Mateo hacia lagunillas, en los alrededores, 17°05’74»S, 65°27’85»W, 600 m, 14 December 2003, I. Jiménez 116 & 119 (AAU!, LPB), Valle de Sajta, 17°08’S, 64°50’W, 220 m, 2 October 2000, M. Kessler et al. 8759, 8827 (UC!LPB), Valle de Sajta, estación experimental de UMSS, 17°05’S, 64°40’W, 288 m, 25 June 1996, T. Killeen 4288 (MO!); Chapare, Territorio Indigena Parque Nacional Isiboro Secure, Cordillera de Mosetenez, 500 m al oeste de la laguna Carachupa, 16°14’S, 66°25’W, 1250 m, 12 September 2007, M. Kessler et al. 13379 (GOET!, LPB!, UC!). La Paz: Charopampa, 500 m, 25 September 2005, R.S. Williams 1336 (UC!); Abel Iturralde, Localidad Candelaria a 6 h de Ixiamas, 13º34’S, 68º40’W, 600 m, 22 April 2001, M.R. Orellana & A. Sajines 1103 (LPB).

Remarks: —Of the species discussed here, Cyathea lasiosora has the widest distribution and as a result shows great morphological variability. Its synonyms represent some morphological extremes.

The type material of Alsophila lasiosora and A. tarapotensis is a collection by Richard Spruce that is stored under different numbers. Most specimens bear either the number “ 4249” or “4349” while few are labelled as “ 4349 (an 4249?)” or “ 4349 (cf 4249)”; some handwritten numbers may also be read as “ 4329”. However, two of these numbers refer unambiguously to angiosperms: Spruce 4329 (S) is a Graffenrieda ( Melastomataceae ) and Spruce 4349 is the type of Hydrangea tarapotensis Briquet (1919: 415) . Only Spruce 4249 is exclusively used for a Cyatheaceae .

Most specimens in London (BM, K) and in Dublin (TCD) have the number 4249, whereas most from the continental herbaria (e.g., B, P) have the number 4349. This transfer error was later imported to the English herbaria through the incorporation of private herbaria (e.g., Herbaria Christensen, Dorfler and Hance). Rosenstock (1909: 291) is the earliest source to cite the ambiguous labelling “ Spruce 4349 (an [sic] 4249?)”, sometimes translated as “ Spruce 4349 (or 4249?)”, and he may have fathered this himself. At P, determination slips by Rosenstock with “ 4349 (cf 4249) ” contradict the original blue labels, which state the collection number to be “ 4249 ” (e.g. P-00631729). Spruce’s annotation of some material with “ 4249 var. videtur ” (which translates as “looks like a variety of 4249 ”) is another indication that Spruce intentionally collected a tree fern under that number, but it also indicates that specimens with this annotation represent a different plant than those labelled only with “ 4249 ”.

The best material for clarifying the situation is deposited at Kew. Here, two sheets have further information about the plants on the blue label typical of Spruce’s collections. The sheet K-000589978 has the information “ Spruce 4249, Tarapoto, at 2500 feet altitude. Caudex 10 feet, slender, naked. Fronds 4–5 feet long, bipinnate. Stipes articulate. (Spruce)”. The material has most pinnules stalked to 3 mm and of linear-lanceolate to weakly triangular shape, with obtuse to acute segments with crenate margins. This morphotype is identical to the two pinnules that constitute B-20- 0000191 (no blue label, “ Peruvia orientalis ,4349 Spruce, com. Godet ”) determined as Alsophila lasiosora Mett. on slips labelled “Herbarium G. Mettenius”. Kuhn (1865: 385) clearly states that he bases the descriptions of the new species solely on specimens and manuscripts left by Mettenius unless otherwise indicated. It is the only specimen found to have been annotated as A. lasiosora by Mettenius himself, and may be regarded as the intended type. With this morphotype further correspond:

- BM-000937696 (Herb. Christensen, no blue label, “ Peruvia orientalis : Tarapoto Spruce 4349 ”)

- BM-000937697 and BM-000937699 (with the original blue labels “ Peruvia orientalis, Spruce 4249 ”, no determination before 1999);

- BM-000937698 (no blue label, Herb. Hance no 6569, “ Spruce 4349”, original determination Alsophila nigra Mart. );

- BM-000937700 (with original blue label annotated with “ prope Tarapoto, Peruviae orientalis , coll. R. Spruce, 1855–6 ” in typical handwriting but “ 4249 var. videtur ” and “ Alsophila tarapotensis Ros. ” apparently added later in different ink and handwriting);

- P-00631726 (Herb. Dorfler, blue label “ Peruvia orientalis, Spruce 4349,” determination slip with“ A. tarapotensis Ros. sp. nov., det. Rosenstock, 1909 ”).

- P-00631727 (blue label with “ Peruvia orientalis, Spruce 4349,”)

The sheet K-000589977 “ 4249 var. videtur ” has the information “ caudex gracilis, 15 pedalis, frondes 4–5 pedalis. Tarapoto, in sylvis rivuli Shilliaco, Majo /56 ”. This and two more sheets at Kew have a “ bis ” added to the number (K-000589975/-000589976), but only the last one also has a blue label. All three sheets represent the same morphotype, which differs only weakly from the type of Alsophila lasiosora in having slightly larger pinnae with shorter-stalked, subsessile and more linear-lanceolate pinnules with blunter, nearly subentire segments. The Kew material includes a small portion of the petiole; the scales and scurf are only preserved in remnants, but are quite typical. The Berlin material B-20-0000190 from the Herbarium G. Mettenius is an entire fertile pinna matching this morphotype, determined as Alsophila nigra Mart. by Mettenius. Besides having an identical labelling as BM-000937698 (which belongs to the other morphotype), it also has a small envelope with the label “ Als. nigra ?, Spruce 4249, Tarapoto ”. Further specimens representing the same plant are:

- P-00631728 (Herb. Dorfler, original blue label with “ 4249 var. videtur ”, paratype of A. tarapotensis );

- P-00631729 (Herb. Dorfler, no blue label, “ 4249 ” on the label by Dorfler, with the “2” written over a rough spot where apparently the “3” had been scratched, and “ 4349 (cf 4249)” on the determination slip from Rosenstock; paratype of A. tarapotensis );

In conclusion, Alsophila lasiosora and A. tarapotensis base on the same collection whose correct number is “ Spruce 4249”. Thus the latter becomes a superfluous name for the former. All samples belong to one species and almost certainly have been taken from one individual plant..Some material on which Alsophila tarapotensis was based came from fronds that have contracted laminar tissue, which is a response to sudden sun exposure of expanding fronds (personal observation). The material originally labelled “ 4249 var. videtur ” can be regarded as a separate collection of the same species, but it only served as paratype for A. tarapotensis .

The type material of Alsophila nigra (Martius s.n.) agrees well with the Spruce material of A. lasiosora at BM (000937696–000937699); it only differs in lacking tortuous hairs on the veins and costules abaxially but having few straight white pluricellular hairs to 1 mm long instead.After my personal observation, tortuous hairs in general but also the white straight hairs of C. lasiosora are deciduous and may be completely lacking in older fronds.

The combination Cyathea nigra Linden ex Fournier (1876: 73) is sometimes used to treat this Neotropical taxon (Forzza et al. 2010), erroneously thinking that it is based on Alsophila nigra Martius (1834: 47) . Fournier described a sterile plant from New Caledonia presented as Cyathea nigra by M. Linden at the “Exposition Internationale de Horticulture de Bruxelles” in 1876. No reference to a type specimen was made but the plant was compared to other New Caledonian species and morphological differences were discussed in detail. As it is an effective publication according to Article 29 of the Melbourne Code ( McNeill et al. 2012), the accompanying color plate may be regarded as the nomenclatural type according to Article 9.1, and the text can be seen as differential diagnosis. Judging from the description, Cyathea nigra may be synonymous with Alsophila stelligera (Holttum 1964: 250) Tryon (1970: 37) but since the plant was sterile and no specimen was preserved, a conclusive assignment to a species will remain impossible. As a matter of fact, the New Caledonian species of Alsophila are all very homogenic regarding scale shape and color. The New Caledonian species of Cyathea sensu stricto can be excluded with certainty because they present a completely different habit of the whole plant (Lehnert 2011).

The synonymous Alsophila kilippii represents a more pubescent form of Cyathea lasiosora . Especially the paratype Krukoff 4937 (BM) has strongly pubescent rhachises, costae and veins. On average, the hairs are mostly longer than in the above-mentioned material of C. lasiosora , persistent and leave a scabrous surface but are still just to 1 mm long. The segment margins are strongly crenate and the sori a bit more distal (i.e., supramedial) than in typeidentical C. lasiosora . This form is frequent in central Amazonia and has probably most apices abruptly reduced (as in the photograph of C. lasiosora in Zuquim et al. 2008). Apparently, this pubescent form is common in the lowlands of the Guayanas while the more typical form of C. lasiosora is found in the highlands and mountains (M. Boudrie, pers. communication). For the time being, A. killipii is considered an eological adaptation of C. lasiosora . Maxon clearly refers to the three sheets of Kilipp & Smith 28133 at US as the type of A. kilipii and to the material deposited at F and NY as the duplicates, which is here interpreted as designation of holotype and isotypes.

The lectototype of the synonymized Alsophila mapirensis from Bolivia has gradually reduced apices and is less hairy than the other types listed in the synonymy, especially contrasting to A. killippii . It comes close to the description of Cyathea atrocastanea , which was compared with C. lasiosora by Labiak & Matos (2009). The depicted material of C. lasiosora corresponds more with the type of A. killipii from Amazonia, and is more contrasting to C. atrocastanea than the type of A. mapirensis and most Andean material of C. lasiosora . Cyathea atrocastanea differs from C. lasiosora in having completely glabrous segments with entire to subentire margins (vs. hairy to glabrescent, margins subentire to crenulate in C. lasiosora ) and having ovate squamules on the costules abaxially (vs. squamules long lanceolate).

Mexia 6270 (MO, W) has typical Cyathea lasiosora- indument but the pinnule outline of C. atrocastanea . It is relatively large and tends in its pinnule shape and size towards C. lockwoodiana but the paraphyses are not thick and conglomerated.

Figures

FIGURE 2. Cyathea spp. in the field: A1–D1. Trunk apex; A2–D2. Lamina apex; showing different expressions of abruptly reduced apices; A3–D3. Details of rhachichses, B–D at junctures with costae, showing presence and extend of pubescence and alae (D3). A. Cyathea pilosissima (Dept. San Martin, Peru). B. Cyathea calamitatis, sp. nov. (Prov. Morona-Santiago, Ecuador). C. Cyathea lasiosora (Dept. San Martin, Peru). D. Cyathea tortuosa (Prov. Zamora-Chinchipe, Ecuador).

FIGURE 3. Cyathea lasiosora. A. Lower petiole, showing scales (Monteagudo et al. 10187, MO). B. Detail of petiole scurf (Monteagudo et al. 10187, MO). C. Junction of rhachis and costa (Monteagudo et al. 10187, MO). D. Pinnule adaxially, glabrescent Andean plant (Monteagudo et al. 10187, MO). E. Pinnule, abaxially, glabrescent Andean plant (Acebey 817, UC). F. Fertile segment, abaxially and adaxially, glabrescent Andean plant (Kessler et al. 13379, GOET). G. Fertile segment, abaxially, hairy Amazonian plant (Cid et al. 3413, MO).