Hypolepis stolonifera var. delasotae Arana, Yañez & Schwartsb., 2014

Arana, Marcelo D., Yañez, Agustina & Schwartsburd, Pedro B., 2014, An updated synopsis of Hypolepis Bernh. (Dennstaedtiaceae) from Argentina, Phytotaxa 188 (2), pp. 91-102 : 93-99

1. Hypolepis stolonifera var. delasotae Arana, Yañez & Schwartsb. , var. nov.

Type: — ARGENTINA. Misiones : Depto. San Pedro. 12 December 1957. R. Capurro 1354 (holotype BA!). Figs. 1 A,B , 2A–H , 4A, B .

Plantae terrestres, inermes, pilosae; rhizomata longe repente; frondibus sparsis, tripinnatis in circumcaesura rhombicis, longe petiolatis, indeterminate crescentia, petiolis rachibusque castaneis; frondulis inferioribus primariis suboppositis, basipedatis, secundariis lanceolatis, tertiariis lanceolatis vel linearis, acutis, crenatis; costulae et laminae infra hirsutis, pilli simplis, strigillosis; sori subglobosi in inferiore latere sinuum laciniarum; sporae monoletae, rotundis leviter reniformibus, papillatis.

A var. stoloniferae differt statura graciliore, segmentis contractis (vs. segmentis applanatis) et laminae infra dense hirsutis (vs. laminae infra glabratis).

Plants terrestrial. Rhizomes long-creeping, branched, bearing several fronds, 2.5–3 mm diam., with catenate acicular hairs, yellowish brown to golden brown, 1.0– 2.5 mm long, 10–20-celled. Fronds erect to arched, 0.2–0.8 m long; petioles sulcate, 10–22(−70) cm long, 2–4 mm diam., inermous, immaculate, glabrescent, with catenate hairs with persistent castaneous basis, sometimes with 1–2-celled, acicular, whitish hairs; laminae rhombic, proximally tripinnate-pinnatifid to quadripinnate-pinnatifid, 22–35 × 30–40 cm; rachises straight, golden brown to light brown, villous, with hairs of two types, catenate, brownish or withish, and unicellular, withish or hyaline, inermous; pinnae strongly ascending; proximal pinnae 8–15 × 13–25 cm, strongly inequilateral, the basiscopic pinnules bigger and more dissected; adaxial side of the costae sulcate, costae abaxially and adaxially villose, the hairs acicular, withish, 0.2–0.4 mm long, 4–8- celled; last pinnules narrowly lanceolate to linear, 2–5 × 8–15 mm, those of the distal pinnae sometimes falcate, usually with acute lobes at the margin and apical lobe acute; costules abaxially villose, adaxially sparsely villose and sulcate, the hairs similar to those from the costae; veins abaxially sparsely villous, the hairs similar to those from the costae and costules but smaller, 0.2–0.4 mm long, 3–5-celled, adaxially glabrous or with very few hairs; laminar tissue between the veins abaxially villous, the hairs similar to those of the veins, adaxially glabrous; lamina margins glabrous; sori marginal, often confluent at maturity; pseudo-indusia conspicuous, proximally green, distally hyaline, membranaceous in the hyaline part, the margin ciliate, the cilia 0.1–0.3 mm long, 3–4-celled; spores yellowish-brown, elliptical in polar view and plane-hemispherical in equatorial view, 31–39 × 22–27 µm, ornamented by baculate processes with a pointed apex, fused together to form crests , and usually connected by branched strands.

Distribution and habitat:— This variety is apparently endemic to “Selva Paranaense” (biogeographic province of Parana Forest according to Morrone 2014), known from the proximities of Río Iguazú, in eastern Misiones ( Argentina) and in western Paraná ( Brazil), inhabiting swampy places. Fig. 6 .

Etymology:— We dedicate this variety to the late Prof. Dr. Elías Ramón de la Sota (1932–2014) in recognition of his exceptional and pioneer work on Neotropical ferns, particularly the Argentinian species of Polypodiaceae and Salviniaceae .

Additional specimens examined (paratypes): — ARGENTINA. Misiones: Depto. Gral. Belgrano, ruta 14, entre Irigoyen y Dos Hermanas, cabecera arroyo campiña de América, en pantano con Dicksonia , 15 November 1970, de la Sota et al. 6146 (BA, LP, fragment RCV); Depto San Antonio, 3 December 1957, Capurro 1275 (BA); Depto. Oberá. 11 September 1945, Capurro 482 (BA) .— BRAZIL. Paraná: Céu Azul, Parque Nacional do Iguaçu, trilha para o Rio Azul , 25º11’20’’S, 53º44’43’’W, 450 m, 4 October 2006, Labiak et al. 3856 (UPCB); Francisco Beltrão, Parque Municipal Irmão Sirilo, 25 October 2006, Schwartsburd et al. 1112 (MBM, SP, UPCB) .

Observations:— This variety differs from Hypolepis stolonifera var. stolonifera by having petioles proximally dark brown and distally straw-coloured (vs. proximally wine-red, medially brown, distally golden brown), shorter laminae 22–35 × 30–40 cm (vs. 60–120 × 80–110 cm), shorter ultimate segments, narrowly lanceolate to linear, usually with acute lobes at the margin and apex (vs. oblong to broadly ovate, with rounded lobes at the margin and apex) and laminar tissue between the veins with abundant acicular pluricellular hairs abaxially (vs. glabrous). The distribution of H. stolonifera var. delasotae is also more restricted, occurring only near the Parana River, whereas H. stolonifera var. stolonifera is found from southern and southeastern Brazil to northeastern Argentina.

Among the north-eastern Argentinian/southern Brazilian Hypolepis species with unarmed petioles and rachises, Hypolepis stolonifera var. delasotae is unique by having rhombic laminae with hairs between the veins abaxially. It further differs from H. poeppigii and H. rugosula subsp. poeppigiana in laminar features and having ciliate pseudoindusia, as shown in the key.

2. Hypolepis stolonifera Fée (1873: 35) var. stolonifera . Type:— BRAZIL. Rio de Janeiro: “Brasilia fluminensi, ad Montes Orgaos, scaturigines amnis Soberbo,” without date, Glaziou 4435 (syntypes: B-200074973!, B-200075126!, C not seen, GH not seen, P-00633493!, P-00633494!, P-00633495!); “Brasilia fluminensi, ad Itatiaia,” Glaziou 5329 (syntypes B-20 0075123!, K!, P-00633496!). Figs. 3A–F , 4C–D .

Rhizomes long-creeping, branched, bearing several fronds, 2–5 mm diam., with catenate, yellowish brown hairs, 1–2 mm long, 15–20-celled. Fronds erect to arching, 1.2–2.0 m long; petioles proximally wine-red, medially brown, distally golden brown, immaculate, glabrescent or with some catenate hairs, inermous, moderately rugose, 45–75(–95) cm long, 3–7 mm diam.; laminae rhombic, proximally tripinnate-pinnatifid to quadripinnate-pinnatifid, 60–120 × 80–110 cm; rachises straight, golden-brown, light brown or greenish, glabrous or with scarce catenate hairs, inermous; proximal pinnae 40–55 × 20–45 cm, strongly inequilateral, the basiscopic pinnules larger and more dissected; pinnae patent; costae abaxially and adaxially glabrescent to sparsely villous, the hairs catenate, hyaline, 0.4–0.8 (–1.2) mm long, 4–8(–10)-celled; last pinnules usually oblong to broadly ovate, with rounded lobes at the margin and apex; costules abaxially villose, adaxially glabrous, the hairs similar to those from the costae; veins abaxially sparsely villose, the hairs similar to those from the costae and costules but smaller, 0.2–0.4 mm long, 3–5-celled, adaxially glabrous; laminar tissue between the veins abaxially and adaxially glabrous; lamina margins glabrous; sori marginal; pseudoindusia conspicuous, proximally green, distally hyaline, membranaceous in the hyaline part, the margins copiously ciliate, the cilia 0.2–0.3 mm long, 3–4-celled; spores yellowish brown, elliptical in polar view and plane-hemispherical in equatorial view, 21–37 × 20–29 µm ornamented by baculate processes fused together to form crests , and usually connected by branched strands.

Iconography: —Fée (1873: tab. 91, fig. 2); Schwartsburd (2012a: figs. 1.A–I, 3A–H);

Distribution and habitat: — Hypolepis stolonifera is the most common species occurring in southern and southeastern Brazil, from the states of Minas Gerais and Rio de Janeiro southwards to Rio Grande do Sul; here it is reported for the first time from Argentina (province of Misiones). It occurs mainly in the mountain regions of the Atlantic forest and forest of Araucaria Jussieu (1789: 413) , especially in forest clearings and at the edges of the forest if it is not too disturbed, in semi-shady places, forming large populations ( Schwartsburd 2012b). Fig. 6 .

Specimens examined: — ARGENTINA. Misiones : Depto. Guaraní. Predio Guaraní, 26º 54’–59’S, 54º 12’–18’W, tramo 1 en camino abandonado, 08 December 2001, Keller 1470 (BA, CTES); Depto. General Belgrano, Parque Provincial Cruce Caballero, sendero Carayá, 26º31’12’’S, 53º59’13’’W, 611 m, 26 November 2013, Zanotti et al. 540 (SI); Depto. San Antonio, 4 December 1957, Capurro 1321 (BA); Depto. San Pedro, Parque Provincial Cruce Caballero, 14 March 2012, Yañez & Marquez 101 (LP) .

3. Hypolepis poeppigii ( Kunze 1834: 50) Rodríguez (1989: 202) . Polypodium poeppigii Kunze (1834: 50) . Hypolepis rugosula (Labill.) J.Sm. var. poeppigii (Kunze) Christensen & Skottsberg (1920: 31) . Phegopteris poeppigii (Kunze) Mettenius (1858: 297) nom. inval. Type:— CHILE. V Región [Región de Valparaíso]: “In turfosis paludosis littoris oceani pacificis prope Concon,” August 1827, Poeppig, Col. Pl. Chil. 264 (holotype: LZ-destroyed; isotypes: B-20 0074807!, BM-001067957!). Fig. 5A, B .

Phegopteris sturmii Philippi (1896: 358) Type:— CHILE. X Región [Región de Los Lagos]: Puyehue, Pilmaiquén, Cueto s.n. (holotype SGO-000000470 image!).

Description and Iconography: — de la Sota (1977 as “ Hypolepis repens ”), Arana & Bianco (2011 as “ Hypolepis repens ”), Schwartsburd (2012a).

Distribution and habitat: —Southern Bolivia (Dept. Chuquisaca), central Chile (regions of Bernardo de O’Higgins (VI), Bío Bío (VIII), Metropolitana de Santiago (XIII) and Valparaíso (V)) and in the northwestern and central Argentina (provinces of Catamarca, Córdoba, Jujuy, Salta, Tucumán). The Sierra de Comechingones in Córdoba represents the southernmost limit of the species in Argentina. It occurs mainly in the mountain regions of the Yungas and Chaco Serrano grasslands, in creek devices and shady places from 800 m above sea level. Fig. 6 .

Observations: —We have selected the correct combination of Polypodium poeppigii into Hypolepis made by Rodriguez (1989 — H. poeppigii (Kunze) R.A. Rodr. ), and not by Maxon (1941 — H. poeppigii (Kunze) Mett. ex Maxon ). Maxon (1941) did not have any intention to do such combination; he only cited “ Hypolepis poeppigii (Kunze) Mett. ”, clearly making confusion between Polypodium poeppigii Kunze and Hypolepis poeppigiana Mettenius (1856) , which are two different entities. At that time, Mettenius (1856, 1858) considered Polypodium poeppigii as Phegopteris poeppigii (Kunze) Mett. , based on Phegopteris Mettenius (1858: 293) , a later homonym of the valid genus Phegopteris ( Presl 1836: 179) Fée (1852: 242) of the family Thelypteridaceae .

4. Hypolepis rugosula ( Labillardière 1806 [1807]: 92) Smith (1846: 8) subsp. poeppigiana (Mett.) Schwartsburd & Prado (2014: 213) . Hypolepis poeppigiana Mettenius (1856: 18) . Type:— CHILE. XIV Región [Región de los Ríos]: “Prope coloniam Arique in prov. Valdivia,” May 1851, W. Lechler, Pl. Chil. 194 (lectotype B 20 0074790!, designated by Schwartsburd & Prado, 2014, isolectotypes B! (2 sheets), FI!, FI-W!, G!, K! (3 sheets), L!, LE!, UPS!, W!). Fig. 5 C–D .

Hypolepis chilensis Fée (1857 [1858]: 76) nom. superfl. for Hypolepis rugulosa based on one share syntype (W. Lechler, Pl. Chil. 194). Syntypes:— CHILE. Valparaiso, [1834 or 1835], Gaudichaud s.n. [31?] (BR!, FI-W!-214906, FIW!-214910, FI-W!-216216, G!, RB!, W!); San Juan Fernandez, s.d., Gay (P?- n.v., PC?- n.v.); XIV Región [Región de los Rios]:”Pres de la colonie Arique, province de Valdivia” [May 1851], W. Lechler, Pl. Chil. 194 (B!-3 sheets [20 0074789, 20 0074790, 20 0074791], FI!, FI-W!-214914, G!-2 sheets, GOET- n.v., K!-000640324, K!-000640325, K!-000640326, L!, LE!-2 sheets, UPS!, W!).

Hypolepis hauman-merckii Hicken (1906: 212) Type:— ARGENTINA. Prov. Buenos Aires: Sierra de la Ventana, 25 December 1905, L.L. Hauman-Merck s.n. (Lectotype SI-000095!, designated by Schwartsburd & Prado, 2014).

Phegopteris poeppigii (Kunze) Fée ex Gay var. hirsuta Philippi (1873: 583) Type:— CHILE. Región XI [Región Aysén del General Carlos Ibañez del Campo]: Puerto Lagunas, January 1872, Simpson s.n. SGO- 000000467 [image!]. (Lectotype designated by Schwartsburd & Prado, 2014).

Description and Iconography: — de la Sota et al. (1998), Schwartsburd & Prado (2014: Fig. 1A, B, E–H ).

Distribution and habitat: — Chile (regions of Aysén del General Carlos Ibáñez del Campo (XI), Araucanía (IX), Biobío (VIII), Coquimbo (IV), Los Ríos (XIV), Magallanes y Antártica Chilena (XII), Valparaíso (VIII) and Juan Fernandez (Más a Tierra and Más Afuera), Chiloé ( Isla Grande). Argentina (provinces of Buenos Aires, Chubut, Mendoza, Neuquén and Río Negro) typically in hyper humid low altitude forests of Patagonia, coasts of lakes and rivers at 0–600 m, also in rock crevices in Pampean Hills. Fig 6 .

Figures

FIGURE 1. Hypolepis stolonifera var. delasotae. A. Habit. B. Abaxial surface of a segment. Scale bar:A = 20 cm; B = 0.3 cm.

FIGURE 2. Hypolepis stolonifera var. delasotae. A. Rhizome long-creeping, branched. B. Hairs of rhizome are catenate, acicular, yellowish brown to golden brown. C. Detail of rachis with two kinds of hairs: hairs catenate, brownish or withish and hairs uni-cellular, withish or hyaline. D. Rachis golden brown to light brown, villose. E. Basal pinnae with strongly inequilateral proximal pinnules. F. Laminar tissue between the veins abaxially villose. G. Detail of last pinnules. Sori marginal often confluent at maturity. H. Pseudo-indusia conspicuous, proximally green, distally hyaline, membranaceous in the hyaline part, the margin ciliate. Scale bars:A = 4 cm; B = 0.3 cm; C = 10 cm; D and E = 0.5 cm; F = 4 cm; G = 2 cm; H and I = 0.2 cm.

FIGURE 3. Hypolepis stolonifera var. stolonifera. A. Rhizome long-creeping, branched. B. Distal Pinna. C. Detail of pinnule. D. Habit. E. Segment with laminar tissue between the veins abaxially glabrous. F. Detail of marginal sori. Pseudo-indusia membranaceous, the margins copiously ciliate. Scale bars:A and B = 5 cm; C = 2 cm; D = 50 cm; E and F = 0.2 cm.

FIGURE 4. A–B. Spores of Hypolepis stolonifera var. delasotae. A. Proximal view at light microscope. B. Proximal view at scanning electron microscope. C–D. Spores of Hypolepis stolonifera var. stolonifera. C. Equatorial view at light microscope. D. Equatorial view at scanning electron microscope. Scale bar:A–D = 10 µm.

FIGURE 5. A–B. Hypolepis poeppigii. A. Habit. B. Detail of segments with rounded lobes at the margin.C–D. Hypolepis rugosula subsp. poeppigiana C. Habit. D. Detail of pinnules abaxially. Scale bars: A and C = 50 cm; B = 2 cm; D = 10 cm.

FIGURE 6. Complete distribution map of Hypolepis poeppigii, H. rugosula subsp. poeppigiana, H. stolonifera var. delasotae and H. stolonifera var. stolonifera.