Salpichlaena papyrus G.G.Cárdenas, Tuomisto & Lehtonen, 2019

Cárdenas, G. G., Lehtonen, S. & Tuomisto, H., 2019, Taxonomy and evolutionary history of the neotropical fern genus Salpichlaena (Blechnaceae), Blumea 64 (1), pp. 1-22 : 11-13

3. Salpichlaena papyrus G.G.Cárdenas, Tuomisto & Lehtonen , sp. nov. — Fig. 3c , 7; Map 1

Salpichlaena papyrus can be recognised by its scales on the abaxial axes having an appearance of crumpled paper, or being broadly ovate with no appendices or only small ones.

— Type: Jones M & Olivas P 165 (holo CR ( CR0241094 ), iso LSCR, TUR (mounted on two sheets: 591746, 591747), USJ), Costa Rica, Heredia, Sarapiquí, La Selva Biological Station , nearest grid post P1000:50 , N10°26' W84°1', 50–150 m, 7 Aug. 2001 .

Etymology. The specific epithet refers to the scales, which have the distinctive appeareance of crumpled paper.

Climbing and non-climbing fronds with herbaceous, chartaceous (to coriaceous) lamina texture, pinnae and pinnule margins not cartilaginous (to slightly cartilaginous); scales on abaxial axes formless resembling crumpled paper or ovate with no appendices or only short ones (1–3 cells); stomata often on white protuberances of the lamina. Non-climbing entire fronds up to 32 by 3 cm. Non-climbing 1-pinnate fronds 20–80 cm. Pinnae up to 9 pairs, apical one lanceolate (elliptic), 11.2–22.2 by 2–2.7 cm; lateral ones lanceolate, parallel-sided (elliptic), 6.3–19.2 by 1.5–2.8 cm; basal ones rarely pinnate; apex acuminate (attenuate), with serrate margin; base obtuse. Climbing fronds with pinnae almost monomorphic, although fertile pinnae usually narrower than sterile. Pinnules subopposite to opposite (alternate). Fertile pinnules in 2–11 pairs; apical one lanceolate, lanceolate-elliptic, elliptic-lanceolate (falcate), 12–27.7 by 0.8–2.1 cm, apex attenuate, serrate margin (to entire), base obtuse, cuneate, equilateral to oblique; lateral ones lanceolate, elliptic, elliptic-lanceolate, parallel-sided (falcate), 12.6–22.7 by 0.6–1.8 cm, apex attenuate, long-attenuate,acuminate, caudate or serrate, base obtuse, equilateral to slightly oblique. Sterile pinnules in 3–10 pairs; apical one lanceolate, parallel-sided (elliptic), 10.7–24.5 by 0.9–3.7 cm, apex attenuate, acuminate, serrate margin, base round, acute, equilateral to oblique; lateral ones lanceolate, parallel-sided (elliptic), 9.6–20.4 by 1.8–3.3 cm, apex acuminate, attenuate (caudate), serrate margin, base obtuse (acute), equilateral to oblique; petiolulate, sessile, rarely winged. Foliar buds rarely present at the base of pinnules.

Distribution — Salpichlaena papyrus occurs in Central America in Costa Rica, Nicaragua and Panama, and in South America in Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru and Venezuela.

Habitat & Ecology — In primary rain forests on variable topographies on relatively well-illuminated sites near paths, tree fall gaps, streams, cliffs and ravines, also along forest borders and in disturbed forests. Reported from clayey to loamy soils. Altitude 50–1830 m.

Additional specimens seen (paratypes). BOLIVIA, Beni, Ballivián, al sud de la Misión Fatima , 950 m, 24 May 1988, Beck G, et al. 16372 ( F). – BRAZIL, Mato Grosso, Chapada dos Guimaraes , Veu das Noivas , 16 Feb.1988, Salino A 402 ( AAU, BHCB, UEC). – COLOMBIA, Meta, Llanos, 457 m, Mar.1948, Sandeman C 5868 ( K); Nariño, 2 km E of Barbacoas, just s of Río Telembí , N1°40' W78°8', 100 m, 13 Oct. 1943, Fosberg FR 21245 ( US). – COSTA RICA, Cartago, forest near the entrance to Parque Nacional Tapanti , 1270 m, 3 Aug. 1983, Moran R 3344 ( AAU, MO); Heredia, Sarapiquí , La Selva biological station , nearest grid post 3400:750 , N10°26' W84°1', 50–150 m, 21 May 2007, Jones M & Putkonen M 610 ( TUR); San Jose, Parque Nacional Braulio Carrillo , Estación La Montura, Sendero A de 0–250 m, 22 Jan. 1984, Gómez LD et al. 20878 ( AAU). – ECUADOR, Morona-Santiago, Mutintz , SE of Makuma , c. km 1 along trail to Tunantza , S2°11' W77°44', 675 m, 31 Oct. 1996, Øllgaard B & Navarrete H 1907 ( AAU); Zamora-Chinchipe, Rd. Zumba-San Andres ,km 1–4, N4°50' W79°8', 1300–1500 m, 24 May 2001, Elgaard Madsen J & Rosales C 8079 ( AAU); Pastaza, Sarayacu, S01°46' W77°29', 12 Apr. 2007, Sirén A et al. 64 ( QCA, TUR). – FRENCH GUIANA, Sommet tabulaire,centre ouest , Tête de crique marécageuse avec éboulis Eclairée et ensoleillée , ± 45km ES de Saül, 600–650 m, Cremers 6502 ( CAY, Z). – GUYANA, Cuyuni-Mazaruni, Mt Maringma,southern slopes of summit escarpment , Arabaru River , N5°12'16.6" W60°34'35.9", 1360 m, 23 June 2004, Clarke et al.12005 ( US); Schomburgk 1140 ( K). – NICARAGUA, Rio San Juan,San Juan del Norte, Reserva Indio-Maiz, N11°07' W83°54', 100 m, 18 Sept.1998, Rueda R et al. 8780 ( US). – PANAMA, Coclé, 3-mountain ridge above El Valle , N8°40' W80°10', 900–1000 m, 24 July 1983, Hamilton C et al. 4121 ( AAU); Colón, Sierra Llorona, collected in forest at P32, N9°21' W79°44', 285 m, 5 Mar. 2008, Jones M & Mitre D 933 ( TUR). – PARAGUAY, in regione calcarea cursus superioris fluminis Apa , Sept. 1913, Hassler 11969 ( K, Z). – PERU, San Martín, Rioja , Sol de Oro, S5°54' W77°27', 1343 m, 17 June 2015, Suominen L et al. 286 ( TUR). – VENEZUELA, Bolivar, Alto Caroni, alrededores de Santa Elena de Uairen , 25 Apr. 1946, Lasser 1642 ( K, US); Zulia , Ayapa , Sierra Perijá, west of Machiques , 1320 m, Dec. 1947, Ginés 137 ( US).

Notes — Salpichlaena papyrus shares the serrate pinnule apices with S. hookeriana , but can be differentiated from it in any ontogenetical state by the scales that are formless and resembling crumpled paper, and in the fertile state by the presence of laminar tissue in the fertile pinnules. Furthermore, the usually higher number of pinnules, more lanceolate pinnule shape and more chartaceous texture of lamina are distinguishing characters.

The scales in S. papyrus are unlike those in any other species with an appearance of crumpled paper without clear shape (more often on the costa), or ovate with no appendices or only small ones (1 or 2 cells) (more often on the rachis and petiolule). Specimens with ovate scales and longer appendices have been observed in Peru, Bolivia and South-East Brazil. Scales on the rachises and petiolules are often big enough to be easily observed by the naked eye.

The serrate pinnule apices of S. papyrus are usually distinctive enough to allow differentiating it from S. volubilis and S. hybrida . However, the serration is less clear in fertile pinnules, which also have thicker lamina texture and more tendency towards a cartilaginous margin than the sterile pinnules do. Therefore, material consisting only of fertile pinnae can be difficult to identify to species, especially if the fronds are already so old that the scales have been eroded.

Foliar buds exist but are very rare in S. papyrus : out of the more than 300 specimens we studied, only two had foliar buds (Øllgaard 105457 AAU and Holm-Nielsen et al. 25341, AAU, both from Ecuador). This contrasts with S. hookeriana , where foliar buds are very common. The non-climbing fronds in S. papyrus are sometimes 2-pinnate, a character that we have not observed in any other taxa.

Salpichlaena papyrus and S. volubilis subsp. thalassica are the only Salpichlaena taxa occuring in Central America and along the western side of the Andes, where they are broadly sympatric. The non-climbing fronds of S. papyrus usually are smaller and have higher number of pinnae than S. volubilis subsp. thalassica .

Figures

Map 1 Distribution of Salpichlaena hookeriana (Kuntze) Alston (■), S. hybrida Lehtonen, G.G.Cárdenas & Tuomisto (▲), S. papyrus G.G.Cárdenas, Tuomisto & Lehtonen (●), S. volubilis (Kaulf.) J.Sm. subsp. volubilis (◆), S. volubilis subsp. amazonica G.G.Cárdenas & Tuomisto (◆), S. volubilis subsp.crenata G.G.Cárdenas & Tuomisto (), S. volubilis subsp.thalassica (Grayum & R.C. Moran) G.G.Cárdenas & Tuomisto (◆).

Fig. 3 Sterile pinnule apices of Salpichlaena. a. S. hookeriana; b. S. hybrida; c. S. papyrus; d. S. volubilis subsp. amazonica; e. S. volubilis subsp. crenata; f. S. volubilis subsp. thalassica; g. S. volubilis subsp. volubilis (a. Tuomisto 12785; b. Christenhusz 2460 3/3; c. Jones 165; d. Tuomisto 16550; e. Christenhusz 2729; f. Tuomisto 15144; g. Christenhusz 4950 2/4; all TUR). — Scale bar: 1 cm.