Physematium jujuiense (Copel.) Ponce & Arana, 2025

Ponce, M. Mónica, Gorrer, Daniel A. & Arana, Marcelo D., 2025, Hidden and neglected taxa inside a collective taxon: taxonomic revision of Woodsiaceae in the Southern Cone of South America, Willdenowia 55 (1), pp. 29-49 : 38-45

2. Physematium jujuiense (Copel.) Ponce & Arana ,

comb. nov. ≡ Woodsia jujuiensis Copel. in Univ. Calif.

Publ. Bot. 19: 297. 1941. – Holotype: Argentina, Jujuy,

Rio Yala, 15 km west of Yala, on half-shady banks, alt.

1900 m, 8 Mar 1936, J. West 6235 (UC [UC 561572!]).

= Woodsia montevidensis var. fuscipes Hieron. in Hedwigia 46: 322. 1907. – Lectotype (designated by Arana & al. 2016: 16): Argentina, Salta, “Prov. de Salta, Los Potreros al pie del Nevado del Castillo, 24.03.1827 ”, P. G. Lorentz & G. H. E. W. Hieronymus 138 (B [B 20 0171577!]; isolectotypes: B [B 20 0171580!, B 20 0171581!], CORD!).

Morphological description — Plants saxicolous, 10– 40 cm tall. Rhizomes suberect, ascending, nodose, compact, scaly, covered by many persistent stipe bases; rhizomatic scales 2–2.8 × 0.4–0.7 mm, triangular-lanceolate, strongly bicolorous, shiny, castaneous to dark brown, with dark brown to blackish central stripe completely sclerotic, margin pale brown, entire or scarcely hairy, apex acute, short or elongate. Fronds fasciculate or approximate. Stipes terete or subterete, brittle, 1/5–1/3 of length of frond, 0.5–1 mm in diam., brown, dark brown to burgundy, blackish at base, basal part with linear-triangular, sclerotic scales, glabrescent, smooth. Blades linear-lanceolate to elliptic-lanceolate, bright green or light green, pinnate-pinnatifid. Rachises subterete to sulcate, stramineous to pale brown, with dense, glandular, uniseriate, pauci-multicellular, articulate hairs. Pinnae numerous, 20–30 pairs, approximate, ascending, triangular, almost 1 cm wide at base, basal pinnae sometimes strongly reduced. Last segments triangular, with margin sparsely incised-crenate, margin hairy, lateral veins visible, free, furcate, tips enlarged to form hydathodes visible adaxially; adaxial and abaxial lamina surfaces with multicellular, uniseriate, articulate hairs, dense on both surfaces, antrorse on adaxial surface. Sori circular, indusium inferior, membranous, with 1–3 striped lobes, usually one of them larger, with marginal, glandular hairs. Spores with folded-reticulate perispore, with complete and incomplete reticules, with some perforations, significant number of spheroids on surface, crest of folds microbaculate. – Fig. 1C, 3A–E, 4A–C, 5A–D.

Distribution — Northwestern Argentina (Catamarca, Jujuy, La Rioja, San Juan, Salta and Tucumán), in Bolivia (Cochabamba) and Peru (Cuzco). This species occurs mainly from the Monte (only in Prepuna district) to Puna biogeographic provinces. – Fig. 7.

Habitat — Physematium jujuiense grows in the high Andean environments, above 2000 m in elevation, on outcrops in rock crevices.

Remarks — This species is very well characterized by its narrowly elliptic blades; stipes thin and brittle, dark brown to almost burgundy, blackish at the bases; and by the rhizomatic and stipe scales linear-lanceolate, shiny and sclerotic.

Additional specimens examined — ARGENTINA: CATAMARCA: Ambato, El Potrero, 15 Mar 1904, Castillón 713 (LIL); Ambato, Rodeo, cerro Lamedero, 1600 m, 12 Feb 1959, V. Carenzo 941 (LIL); Andalgalá, Capillitas, 3000 m, Apr 1944, A. O’Donell 1365 (LIL); Capillitas, Refugio Minero, cerca de arroyo, 3046 m, 13 Jan 2017, J. M. Acosta & S. von Mering 613 (SI); Andalgalá, Río Potrero, 2700 m, H. Sleumer 1845 (LIL); Andalgalá, Rio Pisavil, cerce de El Suncho, 2000 m, 21 Feb 1951, H. Sleumer 1621 (LIL). — JUJUY: Sierra de Zanta, 3500 m, Mar 1931, E. Martin 7513 (LIL); Humahuaca, Ruta Provincial 13, cerca del límite con la provincia de Salta hacia Iruya, 3800 m, 15 Mar 2018, F. O. Zuloaga & al. 16337 (SI); Tumbaya, Volcán, Chilcayo camino a Abra Morada, 2550 m, 26 Feb 1985, R. Kiesling & al. 5708 (SI). — LA RIOJA: Famatina, Sierra de Famatina , camino a la mina La Mexicana, Cuevas de Noroña, 2850 m, 20 Feb 1986, R. Kiesling & al. 6348 (SI); Pelagio Luna, Sierra de Velazco, rancho la Esperanza, 2100 m, Sparre 8650 (LIL); Malanzán, Quebrada de los Nogués, Apr 1940, A. Agüero Vera de Gaudio s.n. (LIL 232851). — SAN JUAN: Angaco, Sierra de Pie de Palo, camino a Mogote de los Corralitos, Aguada del Caño, 2500 m, 15 Feb 1984, R. Kiesling 4406 (SI). SAN LUIS, Pringles, Ruta Provincial 9 entre los Tapiales y La Arenilla, 1600 m, 20 Nov 1984, R. Kiesling & al. 4729 (SI). — SALTA: Orán, Santa Cruz, 3000 m, 20 May 1945, S. Pierotti 1264 (LIL). — TUCUMÁN: Rio Chico, Escaba, cumbre del Marbay, pajonal, 2400 m, L. Monetti 1816 (LIL); Tafí del Valle, La Quebradita, 2200 m, 7 Feb 1959, de la Sota 2076 (LIL); Sierras Calchaquíes, Peñas Azules, 29 Jan 1933, 3400 m, A. Burkart 5536 (SI); Calchaquíes, Quebrada honda, 3500 m, 26 Jan 1952, Sparre 9353 (LIL); Chicligasta, Estancia Las Pavas, Puesto La Cascada, 11 Mar 1924, 2700 m, S. Venturi 3039 (LIL, SI); Trancas, San Pedro de Colalao, 1700 m, 14 Apr 1955, de la Sota 274 (LIL). — BOLIVIA: COCHABAMBA: barrancas del arroyo de Caluya, 3500 m, 20 May 1920, J. Steinbach 4033, 4034 (LIL); Sacaba, Incachaca, sobre rocas húmedas, 2500 m, 14 Oct 1921, J. Steinbach 5862 (LIL). — PERU: CUZCO: Sicuani, Feb 1903, C M. Hicken 3 (SI); Cuzco, 3500 m, Mar 1922, F. Herrera 5 (SI).

3. Physematium montevidense (Spreng.) Shmakov in Turczaninowia 18(2): 12. 2015 [“ montividensis ”] ≡ Dicksonia montevidensis Spreng., Syst. Veg. 4: 122. 1827 ≡ Woodsia montevidensis (Spreng.) Hieron. in Bot. Jahrb. Syst. 22: 363. 1896. – Lectotype (designated by Arana & al. 2016: 15): Uruguay, (“Brasilia”) [Montevideo], Pan d’Açucar, F. Sellow d 517 (B [B 20 0094654!]; isolectotype: B [B 20 0120343!]). = Woodsia incisa Gillies ex Hook. & Grev., Icon.

Filic. 2: t. 191. 1830 ≡ Physematium incisum (Gillies ex Hook. & Grev.) C. Presl, Tent. Pterid. : 66.

1836. – Lectotype (designated by Arana & al. 2016:

15): Argentina, Mendoza, near San Luis, J. Gillies s.n. (BM [BM000937851!]; isolectotypes: BM

[BM000937850!], K [K000229420!]).

Morphological description — Plants saxicolous, 15– 40 cm tall. Rhizomes suberect to decumbent or suberect, separate or spreading, scaly, covered by persistent stipe bases; rhizomatic scales 2–2.8 × 0.4–0.7 mm, ovate-lanceolate or triangular-lanceolate, concolorous, dull, light brown or orange becoming bicolorous, in this case with a central stripe partial to totally sclerotic, brown, margin membranous, broad (twice as wide as central stripe), fimbriate, with spaced hairs, apex ending with an apical hair. Fronds fasciculate. Stipes short to very short, terete or semiterete, 1.5–2 mm long, stramineous, often dark brown proximally, scabrous, with paucicellular, uniseriate, glandular hairs and scaly, scales ovate-lanceolate, dull, concolorous, pale brown, orange or reddish, membranous. Blades triangular, ovate-lanceolate to elliptic-lanceolate, bright green or light green, pinnate-pinnatifid, rarely almost bipinnate. Rachises sulcate, stramineous to pale brown, scabrous or covered by scattered, multicellular, uniseriate, glandular hairs. Pinnae numerous, approximate, triangular, patent to ascending, opposite to alternate, 1–1.5 cm wide at base, basal pinnae gradual to strongly reduced to auriculiform. Last segments subopposite, with acute apex, base adnate, lobes with slightly to strongly crenate margin, sometimes bicrenate, margin hairy, lateral veins visible, free, furcate, tips enlarged to form whitish hydathodes visible adaxially; adaxial and abaxial lamina surfaces with hyaline, multicellular, articulate, glandular, hairs, dense on both surfaces, antrorse on adaxial surface, frequently with yellowish, paucicellular, glandular hairs on abaxial surface. Sori circular, indusium inferior, membranous, saucer-shaped, symmetric, composed of 2 or 3 conspicuous lobes, with glandular hairs on margins. Spores folded-reticulate, with complete and incomplete reticles and perforated surface, also with spinules on crests of folds. – Fig. 1D, 3F–L, 4D–F, 5E–H.

Distribution — Central and northwestern Argentina (Buenos Aires, Córdoba, La Rioja, Mendoza, Río Negro, San Juan, San Luis and Tucumán), southeastern Brazil (Rio de Janeiro, Rio Grande do Sul , Santa Catarina and São Paulo) and Uruguay (Lavalleja, Maldonado, Minas, Rocha and Treinta Tres). This species occurs from the Atlantic biogeographic province, through the Araucaria Forests, Pampean (in isolated mountain systems) to Chaco (only Montane Chacoan district) and Monte biogeographic provinces. – Fig. 8.

Habitat — The species is common in mountainous forests and grasslands, and is found sunny areas, between rocks or in rock crevices, as well as in humid ravines near rivers.

Remarks — Physematium montevidense is distinguished by having rhizomatic and stipe base scales ovate-lanceolate or triangular-lanceolate, concolorous, light brown to orange becoming bicolorous, with centre partially to totally sclerotic, margin broad, with spaced hairs or fimbriate, and apical hair. The stipes are rather robust, especially at the bases, and the laminae are very variable in division, shape and size, depending on the habitats where the plants grow, usually ovate-lanceolate to elliptic-lanceolate, pinnate-pinnatifid to bipinnatifid.

The spore size difference observed in P. montevidense is striking, where our specimens were 26 µm smaller than those analysed by Castro (2004), perhaps because she applied a very broad concept of Woodsia montevidensis . We were able to revise the specimens analysed by her and many of them, identified as “ W. montevidensis ” belong to P. pallidum (large spores) and P. hieronymi (small spores). According to Tryon & Lugardon (1991) Woodsia (= Physematium ) spores with large sizes are related to high ploidy levels. This would indicate that speciation in the genus apparently involves polyploidization.

Additional specimens examined — ARGENTINA: BUENOS AIRES: General Pueyrredón, Estancia La Brava, Sierra Valdez , ladera oeste, 18 Nov 1977, O. Boelcke & al. 806 ( SI) ; Olavarría , Cerro Dos Hermanas, 250 m, 21 Apr 1947, A. Krapovickas 3407 ( LIL) . Tornquist, Cerro Ventana , 7 Dec 1970, A. Burkart & M. E. Múlgura s.n. ( SI 28122 ) ; Coronel Suárez, Camino de Piscultura , 8 Dec 1970, A. Burkart & M. E. Múlgura s.n. ( SI 28123 ) ; Sierra de La Ventana , 23 May 1938, A. L. Cabrera 4470 ( SI) . Saavedra, Pigüé, Cerro Cura Malal , grietas húmedas, 10 Nov 1932, A. Burkart 4667 ( SI) ; Sierra de Curu Malal , abra de la Comenas, 13 Oct 1979, O. Boelcke & al. 91198 ( SI) ; Tandil, Sierras de Tandil, Cerro Leones , 2 Nov 1951, D. Abbiatti 4270 ( SI) . — CÓRDOBA: Calamuchita, Valle de los Reartes, A. Castellanos 197 ( SI) ; Las Guindas, pastizal, 30 Mar 2000, M. Ceballos s.n. (RCVC 3647). Colón, Ascochinga, Tres cascadas, 900 m, 17 Jan 1932, B. Veronesi s.n. ( LIL 436491 ) ; Río Ceballos, Reserva Hídrica La Quebrada, Colanchanga , margen del sendero que une Los Guindos con el río Los Hornillos , 29 Mar 2015, R. E. Morero 414, 415 ( CORD) ; Ascochinga , 28 Nov 1936, E. Nicora 1159 ( SI) ; 28 Nov 1936, M. L. Giardelli 669 ( SI) ; Punilla, Sierra Chica, Cerro Pan de Azúcar , 16 Dec 1886, F. Kurtz 4445 ( CORD, SI) ; Copina , 1400 m, 12 Feb 1947, D. Grassi 2245 ( LIL) ; Los Cocos , 1200, 15 Apr 1962, J. S. Lichtenstein s.n. ( SI 22855 ) , Capilla del Monte , Feb 1933, E. Nicora 150 ( SI) . San Alberto, Pampa de Achala , pastizal, 9 Nov 2000, M. Arana s.n. (RCVC 3727); Pampa de Achala , Ruta provincial 14, proximo al cruce Ruta nacional 20, 1800 km, 9 Jan 2017, J. M. Acosta 590 ( SI) ; San Alberto, Los Gigantes , a 3 km de La Rotonda subiendo por el sendero, 9 Jan 2017, J. M. Acosta 588 ( SI) . San Javier, La Paz – Loma Bola , 21 Apr 1952, J. S. de Lichtenstein s.n. ( SI 18126 ) . Ischilín, Copacabana , 2 Jan 1941, E. Nicora s.n. ( SI 17677 ) . — JUJUY: Valle Grande , entre Tres Morros y Cerro Hermoso, 2940 m, 16 Apr 2016, CMM 792 ( SI) ; Alto Calilegua , 2540 m, 19 Jan 1987, Iudica 387 ( SI) . — LA PAMPA: Depto. Chical Co , 20 Dec 1989, H. O. Troiani 10029 ( SRFA) . Depto. Lihuel Calel, Sierra de Lihuel Calel , 30 Nov 1959, A. Burkart s.n. ( SI 20553 ) ; Lihuel Calel , poco abundante entre grietas a la sombra, 25 Oct 1978, Steibel 5942 ( SRFA) . — LA RIOJA: Cerro Famatina, Guanchin viejo, 25 Jan 1928, Castellanos 28/20 ( LIL 18362 ) ; Velasco , 6 Mar 1944, A. Soriano 981 ( SI) ; cerca de Mina San Juan , 3100 m, Mar 1906, F. Kurtz s.n. ( SI24713 ) . — MENDOZA: San Carlos, Quebrada de Alvarado , 1800–2000 m, 10 Feb 1919, R. Sanzin 3278 ( SI) ; Tunuyán , 25 km al suroeste de Campo de los Andes, 1800 m, J. Araque 1146 ( LIL) ; íbid., M. Cáceres 86 ( LIL) . — RÍO NEGRO: Dpto. San Antonio, Sierra Grande , ladera SE, aprox. 400 m, 8 Feb 1979, S. Crespo & al. 2231 ( BAB) ; ídem, 4 km al sur de Sierra Grande, entre piedras, en Quebrada , 10 Nov 1988, M. Correa & al. 9616 ( BAB) . — SALTA: Ruta provincial 33, Cuesta del Obispo, Paraje La Herradura , 3100 m, 12 Jan 2017, J. M. Acosta 606 ( SI) . — SAN JUAN: Valle Fértil, Sierra de Elizondo , 1200 m, 16 Nov 1987, M. E. Múlgura & al. 804 ( SI) . — SAN LUIS: Cerro el Morro, 14 Oct 1949, Castellanos s.n. ( LIL 380139 ) ; Merlo, Peñón Colorado , 19 Feb 1936, M. A. Vignati 37 ( SI) ; Juan Martín de Pueyrredón , El Volcán, 25 Jan 1908, F. Pastore 13 ( SI) ; Coronel Pringles, Ruta Provincial 9, Junin , Piedra Blanca , ladera occidental de Comechingones, 2000 m, D. Grassi 2166, 2171 ( LIL) ; La Carolina, Monumento – Museo Lafinur, 1680 m, 28 Nov 2008, F. Biganzoli, R. León & C. Larsen 2037 ( SI) ; Dique La Florida , cerca del vertedero, 15 Dec 1991, M. E. Múlgura 1173 ( SI) ; Estancia Grande , 28 Feb 1939; M. A. Vignati 7063 ( SI) . — TUCUMÁN: Tafí, Valle de Tafí , Mar 1908, C. Bruch s.n. ( SI 24691 ) ; Tafí, San José , 2100 m, 17 Feb 1949, R. Sparre 5873 ( LIL) , Los Chamicos , 900 m, 30 Jan 1924, S. Venturi 2778 ( SI) . — BRAZIL: RIO DE JANEIRO: Nova Friburgo, Parque Estadual dos Três Picos , na subida para caixa de Fósforo, 22°24'00"S, 42°44'00"W, 1550 m, Jun 2006, J. Condack 442 ( RB) . — RIO GRANDE DO SUL : Soledade, Sep 1913, C. Jürgens 355 (NY 678816); Santa Anna, 26 May 1907, W. G. F. Herter 3097 (NY 678817); Jaquirana , Parque Estadual do Tainhas. Estrada para o passo do S, 18 Dec 2021, F. Gonzatti & al. 6219 ( HUCS) . — SANTA CATARINA: São Joaquim , ad saxa aprica juxta flum. S. Matheus, 1907, L. Spannagel s.n. (Herb. Rosenstock 381) (P01402552, SI) ; idem, Bom Jardim da Serra , entre pedras no campo, 1300 m, 15 Dec 1958, R. Reitz & R. M. Klein 7902 ( PACA) ; Lages, Campo Bello, 27°48'58"S, 50°19'33"W, sine col. (NY01016492); Campos dos Padres , em barrancos, 1800 m, 19 Dec 1948, R. Reitz 2622 ( RB) . — SÃO PAULO: Campos do Jordão , 20 Feb 1937, P. Campos Porto 3111 ( RB) . — URUGUAY: Uruguai, Banda Oriental Del Uruguay , 1 Jan 1816, A. Saint-Hilaire C2/2148 (P) . — FLORIDA: Cerro Colorado, estancia San Pedro , 2 Oct 1942, Gallinal, Aragone, Bergalli, Campal & Rosenhairs, 2-multicellular, articulate. Sori circular, few per segment, indusium inferior, membranous, central, bowl-shaped, somewhat globose, divided into 2 or 3 lobes, with glandular hairs on margin. Spores with folded-reticulate perispore, with complete and incomplete reticules, with rugulatescabrate surface; margin of folds echinulate. – Fig. 1B, 3M–R, 4G–I, 5I–L .

gurtt 5030 (LIL). — LAVALLEJA: en intersticio rocoso, 14 Dec 1948, Rosengurtt 5318 ( LIL) ; cerro de Arequipa, Apr 1907, A. Lombardo s.n. ( LIL 225539 ) ; Sierra de la Ballena, 4 Aug 1940, Legrand 2229 ( LIL) ; 12 Jun 1985, Del Puerto, Davies & Berrutti 17847 ( MVFA) . — MALDONADO: Sierra de las Ánimas , 500 m, 3 May 1931, W. Herter 885a ( LIL) ; Sierra de las Ánimas , 6 Mar 1915, M. B. Berro s.n. ( MFVA) ; en lugar húmedo y soleado, E. Marchesi 1359 ( MFVA) ; Sierra de las Ánimas, Cerro Betete , entre piedras, a la sombra, 27 Oct 1996, M. Bonifacino & E. Mendez s.n. (MFVA 25864); Cerro Lagunitas entre piedras, 1 Apr 1999, M. Bonifacino s.n. (MFVA 28870). — MINAS: cerro Verdún, 4 Dec 1900, M. B. Berro 1420 ( MFVA) ; — ROCHA: Cerro de San Miguel , A. Castellanos 17451 ( LIL) . — TREINTA Y TRES: Yerbal , 200 m, Apr 1928, G. Herter 885 ( SI) .