Pteris arachnoidea Kaulfuss (1824: 190)
Allosorus arachnoideus (Kaulf.) Presl (1836: 153)
Allosorus arachnoideus (Desv.) C.Presl
Pteris aquilina
var. arachnoidea (Kaulf.) Eaton (1861: 203)
Cincinalis arachnoidea (Kaulf.) Trevisan de Saint-Léon (1874: 31)
Pteridium aquilinum
var. esculentum (G.Forst.) Kuhn (1882: 27) f. arachnoideum (Kaulf.) Hieronymus (1909: 246)
Pteridium aquilinum
var. arachnoideum (Kaulf.) Brade (1920: 56)
Pteridium arachnoideum (Kaulf.) Maxon (1924: 89)
aquilina
Farwell (1931: 290)
var. arachnoidea (Kaulf.) Farwell (1931: 290)
Pteridium caudatum
subsp. arachnoideum (Kaulf.) Lellinger (2003: 150)
Schwartsburd et al. 2014: 110
Aspidium brasilianum Presl (1822: 176)
Cystopteris brasiliana (C.Presl) Presl (1836: 93)
Hypolepis brasiliana (C.Presl) Kuhn (1882: 347)
Pteris psittacina Presl (1822: 185)
Allosorus psittacinu
Presl (1836: 153)
Allosorus psitaccinus
Pteris aquilina
var. psittacina (C.Presl) Baker (1870: 403)
Pteridium psittacinum (C.Presl) Maxon (1933: 141)
Pteridium aquilinum
subsp. psittacinum (C.Presl) Christensen (1934: 166)
Cheilanthes aquilinaris Fée (1873: 37
Hypolepis aquilinaris (Fée) Christ (1901: 636)
Schwartsburd et al. 2014: 110
Formal recognition of six subordinate taxa within the South American bracken fern, Pteridium esculentum (P. esculentum subsp. arachnoideum s. l. - Dennstaedtiaceae), based on morphology and geography
Schwartsburd, Pedro B.
Yañez, Agustina
Prado, Jefferson
Phytotaxa
2018
2018-01-05
333
1
22
40
5KRMK
(Kaulf.) Thomson (2012: 45)
Thomson (2012: 45) sensu Thomson
45
2012
Kaulf.
[164,1241,1760,1787]
Polypodiopsida
Dennstaedtiaceae
Pteridium
Plantae
Polypodiales
5
27
Tracheophyta
subSpecies
stricto
esculentum
arachnoideum
Two varieties are recognized:
1. Margins of sterile laminae regularly transformed into continuous pseudo-indusia; simple distal segments linear, to 3(–4) cm long ............................................................................................................................................................................... var. arachnoideum -. Margins of sterile laminae mostly not transformed into pseudo-indusia and ciliate; simple distal segments triangular elongate, to 1.5 cmlong .......................................................................................................................................................... var. paedomorficum FIGURE 3.A–C. Pteridium esculentumsubsp. gryphusvar. gryphus: A. pinnule ( Gleason 423), B. segment, abaxially ( Gleason 423), C. segment, cross section, abaxial side up, showing sericeous veins, with lax, arachnoid hairs, and laminar tissue between the veins with farinaceous appearance, fully covered by gnarled hairs ( Gleason 423), D–F. Pteridium esculentumsubsp. gryphusvar. harpianum: D. pinnule (“ B.W.” 1121[US-2019769]), E. segment, abaxially ( Barreto 1623), F. segment, cross section, abaxial side up, showing veins abaxially nearly glabrous, and laminar tissue between the veins with farinaceous appearance, fully covered by gnarled hairs ( Barreto 1623). Drawn by R. Pinto. 1a. Pteridium esculentum(G.Forst.) Cockayne subsp. arachnoideum(Kaulf.) Thomson var. arachnoideum( Figs. 1D–F, 2, 6, 7)
Pteris arachnoideaKaulfuss (1824: 190), nom. cons. Allosorus arachnoideus(Kaulf.) Presl (1836: 153)[as Allosorus arachnoideus(Desv.) C.Presl]. Pteris aquilinaL. var. arachnoidea(Kaulf.) Eaton (1861: 203). Cincinalis arachnoidea(Kaulf.) Trevisan de Saint-Léon (1874: 31). Pteridium aquilinum(L.) Kuhn var. esculentum(G.Forst.) Kuhn (1882: 27)f. arachnoideum(Kaulf.) Hieronymus (1909: 246). Pteridium aquilinum(L.) Kuhn var. arachnoideum(Kaulf.) Brade (1920: 56). Pteridium arachnoideum(Kaulf.) Maxon (1924: 89). Filix-foemina aquilina(L.) Farwell (1931: 290) var. arachnoidea(Kaulf.) Farwell (1931: 290). Pteridium caudatum(L.) Maxon subsp. arachnoideum(Kaulf.) Lellinger (2003: 150). Lectotype(designated by Schwartsburd et al.2014: 110):— BRAZIL. State unknown: “Habitat in Brasilia”, without date, C.A. de Chamisso s.n.(LE!). = Aspidium brasilianum Presl (1822: 176), nom. rej. Cystopteris brasiliana(C.Presl) Presl (1836: 93). Hypolepis brasiliana(C.Presl) Kuhn (1882: 347). Lectotype(designated by Schwartsburd & Prado 2011):— BRAZIL. Rio de Janeiro: “ Ad Rio JaneiroBrasilia legit def.” [In via ad S. João Marcos ad Engenho da Varge, Capit. Rio de Janeiro], [ March 1818], J.B.E. Pohl s.n.[ 3794] ( lectotype PRC!, isolectotypeW!). = Pteris psittacina Presl (1822: 185), nom. rej. Allosorus psittacinus (C.Presl) Presl (1836: 153)[as Allosorus psitaccinus]. Pteris aquilinaL. var. psittacina(C.Presl) Baker (1870: 403). Pteridium psittacinum(C.Presl) Maxon (1933: 141). Pteridium aquilinum(L.) Kuhn subsp. psittacinum(C.Presl) Christensen (1934: 166). Lectotype(designated by Schwartsburd & Prado 2011):— BRAZIL. Rio de Janeiro: “ Ad Rio JaneiroBrasilia” [Circa Rio de Janeiroin collibus], [ December 1818], J.B.E. Pohl s.n.( lectotype PRC!, isolectotypeW!). = Cheilanthes aquilinaris Fée (1873: 37, t. 91, fig. 1). Hypolepis aquilinaris(Fée) Christ (1901: 636). Lectotype(designated by Schwartsburd et al.2014: 110):— BRAZIL. Rio de Janeiro: “Brasilia fluminensis”, [Haut Itatiaia], [ 6 June 1871], A.F.A. Glaziou 5330( lectotypeB!-200074529, isolectotypesC n.v., HBG n.v., K!-000640338, NY-fragment [00144358, image!], P-00633378 [ex Herb. Fée, image!], P-00633379 [image!], P-00633380 [ex Herb. Glaziou, image!], P-00633381 [ex Herb. Glaziou, image!], S-05- 9822 [image!], S-10-34455 [image!], US-00066403 [image!]). Plantsforming extensive thickets. Rhizomeslong-creeping. Frondsto 3(–4.5) m long; petiolesproximally with epipetiolar roots; laminaeproximally 3–4-pinnate-pinnatifid; pinnae and pinnulesdistally with free lobes between the segments; compound distal segmentsinequilateral, irregularly dissected, caudate at apex; simple distal segmentslinear, to 3(–4) cm long; costaeabaxially with reddish catenate hairs, 5–7-celled, 0.3–0.5 mmlong, adaxially glabrous; costulesabaxially with reddish catenate hairs, adaxially glabrous; veinsabaxially sericeous, with lax, arachnoid hairs, 0.6–0.8 mmlong, adaxially glabrous; laminar tissue between the veinsabaxially glabrous and totally visible, without farinaceous appearance, adaxially glabrous; laminar marginsregularly transformed into pseudo-indusia; pseudo-indusiaglabrous.
Distribution and ecology:— Colombia, Venezuela, Guyana, French Guiana, Peru, Bolivia, Paraguay, Brazil, Uruguay, and Argentina; possibly also in Suriname, Central America and Mexico; from sea level to 2200 m( Figs. 2, 6, 7). Additional specimens examined(see also Schwartsburd et al.2014for additional specimens from eastern South America):— COLOMBIA. Cesár: Sierra de Perijá, Manaure, 1585 m, 31 January 1945, M. L. Grant10719(US). Magdalena: Mount Auyantepui, 1200–1800 m, 21 May 1937, F. Cardona62( US).— VENEZUELA. Distrito Capital: Southslope of Cerro Avila, 27 December 1939, C. Vogl59( LIL).— GUYANA. Takutu Superior: Kanuku Mountains, 975 m, 22 April 1938, A. C. Smith3657( NYn.v., US).— FRENCH GUIANA. St. Laurent du Maroni: Massif Dékou-Dékou4, 4°42’N, 53°57’W, 480 m, 24 November 2000, J. J. de Granvilleet al. 14388( CAYn.v., Pn.v., US).— PERU. Localityunknown [Yamatiliá?], November 1920, C. Bues1356 (US); Localityunknown [ Yamapaca], 1700 m, November 1932, C. Bues1999 (US).— BOLIVIA. Beni: Ballivian Province, 25 kmfrom Yucumo, 15°17’S, 67°04’W, 950 m, 16 July 1990, A. Fay& L. Fay2723( MO n.v.,US). La Paz: Charopampa, 1901–1902, R. S. Williams1183( US); Nord Yungas, Polo-Polobei Coroico, 1100 m, October–November 1912, O. Buchtien3542( US); Nord Yungas, Millunguaya, 1300 m, December 1917, O. Buchtien, Herb. Boliv.796( US); Sud Yungas, 2100 m, 3 December 1906, O. Buchtien455( US). Santa Cruz: Florida, 18°14’23”S, 63°40’40”W, 1520 m, 18 January 2006, M. Nee& J. Wen53833( US); San Carlos Mapin, 750 m, October 1907, O. Buchtien1133(US). Provinceunknown: Withoutlocality, 1851, Weddells.n.( US).— BRAZIL. Bahia: Chapada Diamantina, 11°35’28”S, 41°12’27”W, 15 March 2008, M. L. Guedeset al. 14217(US); Santa Cruz de Cabrália, 16°23’S, 39°08’W, 80–100 m, 5 July 1979, S. A. Mori et al. 12105( US). Ceará:Near Guaramiranga, 680 m, 22 August 1944, H. C. Cutler 8241( US). Distrito Federal: Chapada da Contagem, 700–1000 m, 19 August 1964, H. S. Irwin& T. R. Soderstrom5301(US-on 2 sheets). Espírito Santo: Linhares, Comboios, 19 September 1991, L. Behar& G. M. F. Viégas354( VIES); Marechal Floriano, 20.4131°S, 40.6831°W, 700 m, 12 August 1988, O. J. Pereira& L. Behar1629( VIES); Santa Maria de Jetibá, 20°08’S, 40°49’W, 6 September 2008, T. S. Lorenciniet al. 49( VIES); Santa Teresa, 17 May 1985, W. Boone452( MBML). Goiás: Chapadados Veadeiros, 14°S, 47°W, 1000 m, 14 February 1966, H. S. Irwinet al. 12817(US-on 2 sheets); Goiânia, entre Goiâniae Nerópolis, 4 July 1968, J. A. Rizzoet al. 1671( UFG); Serra dos Pirineus, 1230m, 16 May 1973, W. R. Anderson10386( US); Cocalzinho, Serra dos Pirineus, 17°47’56”S, 48°49’07”W, 1150 m, 18 August 2007, P. G. Delprete et al. 10293( RB). Mato Grosso do Sul: Ponta Porã, Fazenda Curupi, 20 November 1999, I. A. Carneiro 1( SP). Minas Gerais:Fazenda São José, 12 September 1963, R. R. Santos28653( LP). Paraná: Ponta Grossa, 25°08’27”S, 50°01’46”W, 1000 m, J. Prado& R. Y. Hirai2351( SP); Tijucas do Sul, 14 February 1978, A. Krapovickas& C. L. Cristobal33652( LP). Rio de Janeiro: Cachoeira de Macacu, Boa Vista, Santa Fé, 22°27’S, 42°37’W, 4 July 2014, P. B. Schwartsburd et al. 3323( VIC). Rio Grande do Norte:São Bento, 6°25’28”S, 35°41’57”W, 24 August 2012, J. G. Jardim et al. 6383( RB). Rio Grande do Sul:Caxias do Sul, 5 March 1949, Afonso s.n.( LP). Roraima: Serra dos Surucucus, 2°42– 47’ N, 63° 33–36’ W, 18 February 1969, G. T. Prance et al. 10059( An.v., Kn.v., INPA n.v., MGn.v., NYn.v., Rn.v., Sn.v., US-on 2 sheets). Santa Catarina: Florianópolis, Morro da Cruz, 30 July 2000, 120 m, D. B. Falkenberg 2000( SP); Monte Castelo, Lajeadinho, 23 February 2006, G. E. Barboza1610( CORD). São Paulo: Pindamonhangaba, Pico do Itapeva, 22°46’S, 45°32’W, alt. 1888 m, J. Prado& R. Y. Hirai2337( SP, US).— PARAGUAY. Caaguazú: Est. Itá Carú, 29 December 1972, A. Schinini5826( MO). Caazapá: Tavai, 26º10’S, 55º17’W, 21 December 1988, N. Soria3020( MO). Cordillera: Cerro Ybitú Silla, 1 km Sof Tobatí, 25º 12’ S, 57º 07’ W, 28 May 1988, E. Zardini4398( MO). Paraguarí:National Park Ibicu’í, Road to Cesar Barrientos, 26º03’ S, 56º 50’W, 15 September 1988, E. Zardini 7237( MO).— URUGUAY. Maldonado: Pan de Azucar, March 1947, A. Lombardo s.n.( LIL). San José: Sierra de Mahona, 18 October 1941, Rosengurtt B-3521(US). Tacuarembó:Gruta de los cuervos, 31°37’19.1”S, 56°02’34.7”W, 194 m, 28 February 2012, E. L. Meza Torres et al. 1447( RB).— ARGENTINA. Buenos Aires:General Pueyrredón, Sierra La Peregrina, 13 December 1982, Alo& Mon842( BAL); Cerro Copelina, 29 January 1932, C. M. Hicken512( SI). Catamarca:El Rodeo, 1500 m, 25 November 1910, Castillón s.n.(US-1113401); El Rodeo, 27 May 1910, Castillóns.n.(US-2084229). Córdoba: Depto. Calamuchita, Sierra Grande, Cerro Blanco, 26 August 1952, A. T. Hunziker9472( CORD); San Alberto, Sierra Grande, Mina Clavero, s.d., Burkart7757( SI); Depto. San Javier, Los Molles, 3 September 2005, R. E. Morero120( CORD); Sierra de Achala, 15 September 1889, F. Kurtz6538( SI). Corrientes: Mburucuyá, 25 June 1952 T. M. Petersen1761(US-on 2 sheets); Berón de Astrada, January 1977, A. Schinini14126( LP). Dept. General San Martín, Tres Cerros, 8 November 1936, A. Burkart8030( SI); Dept. San Miguel, 12 km NEof San Miguel, Ea. Curuzú Laurel, 27 April 1975, A. Schininiet al. 11441( MO, SI). Jujuy: Termas de Reyes, cerros, 17 March 1940, A. Burkart& N. S. Troncoso11235( SI); Depto. Manuel Belgrano, Sierra de Zapala, 14 November 2002, F. O. Zuloagaet al.7614( SI). Misiones: Dept. Candelaria, Santa Ana, 19 September 1912, Rodriguez627( SI); Dept. General Manuel Belgrano, 8 kmfrom Bernardo de Irigoyen, 22 May 1991, Delucchi519(LP-on 2 sheets); Dept. Guaraní, Predio Guaraní, 15 October 1997, S. G. Tressens5934( SI); Dept. Iguazú, Puerto Wanda, 16 December 1950, J. E. Montes10410( LP); Dep. San Ignacio, Teyucuaré, 15 July 1945, A. Krapovickas2421( LP); Dept. San Pedro,borde de Ruta Provincial14, 21 February 2016, A. Yañez& G. Marquez167( LP); Piñalito Provincial Park, 17 February 2016, A. Yañez& G. Marquez142( LP). Salta: Caminoa Jujuy, Caldera, 1 February 1949, A. L. Babrera& H. Schwabe223( US). Tucumán: Famaillá, La Fronterita, 500 m, 19 February 1922, S. Venturi1721( US). Notes:— Pteridium esculentumsubsp. arachnoideumvar. arachnoideumis the most common morphotype in eastern and southern South America, but also reaches other regions ( Figs. 2, 6, 7). It is characterized by free lobes between the ultimate segments, inequilateral, irregularly dissected compound distal segments, which are caudate at apex, simple distal segments to 3(–4) cm long, lax, arachnoid hairs abaxially on veins, giving a sericeous appearance, and laminar tissue between the veins abaxially glabrous, without farinaceous appearance (gnarled hairs absent) ( Figs. 1D–F). 1b. Pteridium esculentum(G.Forst.) Cockayne subsp. arachnoideum(Kaulf.) Thomson var. paedomorficumSchwartsb. & J.Prado, nom. nov.( Figs. 5A–E, 6) Pteridium aquilinum(L.) Kuhn var. umbrosumChrist ex Rosenstock (1907: 90)[as Pteridium aquilinumvar. umbrosa], nom. illeg.(non Luerssen 1889: 107). Lectotype(designated here).— BRAZIL. Rio Grande do Sul: Rio Pardo, Fazenda Soledade, 1906, L.C. Jürgens & A. Stier 118(NY!, probable isolectotypesMVM! [“ Jürgens & Stier 190 b”], SI! [“ Jürgens 267”], US! [“ Jürgens 267”]). Plantssparse, not forming thickets. Rhizomesshort-creeping. Frondsto 1.5 mlong; petiolesproximally with epipetiolar roots; laminaeproximally 3-pinnate-pinnatifid; pinnae and pinnulesdistally occasionally with free lobes between the segments, generally with only a few decurrent lobes; compound distal segmentsequilateral, regularly dissected, not caudate at apex; simple distal segmentstriangular elongate, to 1.5 cmlong; costaeabaxially with reddish catenate hairs, 5–7-celled, 0.3–0.5 mmlong, or sometimes glabrous, adaxially glabrous; costulesabaxially with reddish catenate hairs, or sometimes glabrous, adaxially glabrous; veinsabaxially with setose to arachnoid hairs, 0.3–0.6 mmlong, adaxially with a few setose hairs or glabrescent; laminar tissue between the veinsabaxially glabrous and totally visible, without farinaceous appearance, adaxially glabrous; laminar marginsirregularly transformed into pseudo-indusia, copiously ciliate; pseudo-indusiainterrupted, copiously ciliate. Distribution and ecology:—Possibly the same as Pteridium esculentumsubsp. arachnoideumvar. arachnoideum ,with definitive records from Brazil, Bolivia, Paraguay, Uruguay, and Argentina( Fig. 6).
Etymology:—The varietal epithet is an allusion to the pedomorphic nature of this taxon. Specimens examined:— BOLIVIA. La Paz: Charopampa, Mapini, 570 m, November 1907, O. Buchtien1135(US).— BRAZIL. Minas Gerais: Alto Caparaó, P. N. Caparaó, 29/IV/1989, A. Salinoet al. s.n.(UEC-57698); Bandeira, Mata do Boi Rajado, 15°48’18”S, 40°30’58”W, 662 m, 1 March 2004, A. Salinoet al. 9406( BHCB); Caldas, 5 September 1873, H. Mosén, ex Herb. Bras. Regnell.2065( BM, K, LE); Conceição do Mato Dentro, 19°06’03.1”S, 43°34’26.9”W, 1250 m, 13 September 2003, A. Salinoet al. 8938( BHCB); Diamantina, Trinta-Réis, 13 March 1989, R. F. Novelino631(BHCB- n.v., CESJ, SPF); Moeda, Serra da Moeda, 5 October 1985, M. J. Chagas& T. S. M. Grandi6614( UEC); Moeda, Serra da Moeda, 20°19’22”S, 43°56’30”W, 1277 m, 7 September 2006, G. Heringer91( BHCB); Ouro Preto, P. E. Itacolomi, 13 May 1998, A. Salinoet al. 4240( BHCB); Rio Acima, RPPN Andaime, 18 February 2004, J. B. Figueiredo& J. C. Queiroz254( BHCB); Santa Maria do Salto, Distrito Talismã, 16°24’16.5”S, 40°03’27.4”W, 750–850 m, 10 October 2003, A. Salinoet al. 9238( BHCB); São Thomé das Letras, Camping do Noel, 7 June 2009, P. B. Schwartsburdet al. 1997( SP); Uberlândia, Estação Ecológica do Panga(bark spore bank experiment), 12 July 1997, M. A. Ranal776( HUFU n.v., SP); id., 12 July 1999, M. A. Ranal654( SP); id., 31 July 1999, M. A. Ranal793( SP); id., 28 November 1999, M. A. Ranal991( SP). Paraná: Bateias, 20 April 2005, P. B. Schwartsburdet al. s.n.(UPCB-54124); Bateias, Alto da Serra, 24 April 2005, P. B. Schwartsburdet al. 763( MBM); Campina Grande do Sul, Morro Capivari Grande, 1400 m, 4 March 2007, P. B. Schwartsburd& A. M. X. Lima1247( SP); Guaporema, 21 July 1988, M. V. K. Endos.n.(MBM-124178); Indianópolis, 21 July 1988, M. V. K. Endos.n.(MBM-124179); Nova Olímpia, 23 April 1988, J. C. S. Pereiras.n.(MBM-124174); Piraquara, Pinhais, 1 March 1970, G. Hatschbach23987( MBM); Planalto, 17 August 1988, J. J. Vituris.n.(MBM-133358 p.p.); Quatro Barras, Graciosa, 27 January 1988, M. C. Antuness.n.(MBM-124182); Rondon, Indianópolis, 25 February 1988, M. V. K. Endos.n.(MBM-124191); Santo Antônio do Sudoeste, Cerro Negro, 31 August 1988, C. Alberto Bonezzis.n.(MBM-133366); São Jorge D’Oeste, 17 August 1988, M. S. Matzenbachers.n.(MBM-133368); São Mateus do Sul, BR-476, km 152, 26 January 1985, A. Krapovickas& C. L. Cristobal39667( LP); São Miguel, Dois Vizinhos, 7 January 1988, M. A. Mascarenhass.n.(MBM-124183). Rio de Janeiro: Cachoeira de Macacu, Boa Vista, Santa Fé, 22°27’S, 42°37’W, 4 July 2014, P. B. Schwartsburdet al. 3328( VIC); Itatiaia, P. N. Itatiaia, estr. p/ Abrigo Rebouças, km 12, 04 November 1965, R. M. Tryon& A. F. Tryon6734(HB-on 2 sheets); Localityunknown, ded. 1838, Anonymous[unreadable] s.n.( B). Rio Grande do Sul: Santa Cruz, Fazenda Tangerina, s.d., L. C. Jürgens& A. Stier95( S?- n.v.[ Syntypeof P. aquilinumvar. umbrosum). Santa Catarina: Hammonia, October 1911, H. Luederwaldts.n.(NY-00815815, image!, SP-21261); Hammonia, October 1911, H. Luederwaldts.n.(NY-00815816, image!). São Paulo: São Paulo, Ipiranga, August 1907, H. Luederwaldts.n. (SP-21265); Municipalityunknown (probably near Santo André), Estação Alto da Serra, June 1910, H. Luederwaldts.n.(SP-21266 [on 2 sheets]); Municipalityunknown, s.d., Anonymous[unreadable] s.n., Rosenstock Filic.Austro- Bras.s.n.( US). Stateunknown: Localityunknown, s.d., W. J. Burchell 1076( B, K, LE); Localityunknown, s.d., W. J. Burchell 1401( B, K); Localityunknown, 11 September 1948, A. Macedo1218- A( BM).— PARAGUAY. Caaguanzú: 22 km Sde Caanguazú, Estrada Itá-Carú, 29 December 1972, A. Schinini5826( MBM). Paraguarí: Parque Nacional Ybycuí, 26º03’S, 56º60’W, 23 June 1988, E. Zardini4942( MO); Dep. Misiones, Isla San Pablo, 6 April 1988, S. Keel V-64( LP).— URUGUAY. Artigas:Catalan, November 1927, Herter 82683( SI). ARGENTINA. Buenos Aires: Mar del Plata, 6 January 1933, C. M. Hicken s.n.(SI-07232). Chaco:Puente Río Canghí Chico, 28 January 1952, Meyer17228( MO); Fontana, March 1928, T. Meyer39( SI). Cordoba: Caminoa Villa Benegas, 29 November 2016, Gutierrezet al. 1133( BA); Depto. San Javier, San Javier, s.d., Castellanos s.n.(BA-10089); Los Molles, s.d., Castellanos.n.(BA-10066 p.p.). Corrientes: Dept. Mburucuyá, Estancia Santa Teresa, s.d., T. M. Petersen1761( LP); Dept. San Martín, Tres Cerros, Cerro de Susini, 16 September 1979, A. Schininiet al. 18552( LP). Misiones: Dept. Cainguás, Puerto Rico, 16 May 1949, J. E. Montes3892( LP); Colonia2 de Mayo, 22 May 1951, A. L. Cabreraet al. 335( LP); Dept. Iguazú, 20 July 1918, G. T. Bertoni1019( SI); Dept. Montecarlo, Puerto Piray, Alto Paraná, 21 October 1901, T. Stuckert11509( CORD); Dept. San Ignacio, Teyú Cuaré, 11 July 1937, Pérez Moreaus.n.( BA); San Pedro, Esmeralda Provincial Park, 2016, A. Yañez& G. Marquez143( LP). Tucumán: Famaillá, La Fronterita, November 1922, S. Venturi1721( BA); Valle de Tafí, December 1911, L. Castillón48( SI). Notes:— Pteridium esculentumsubsp. arachnoideumvar. paedomorficumis a fully fertile, pedomorphic form of P. esculentumsubsp. arachnoideum ,occurring in the same areas of P. esculentumsubsp. arachnoideumvar. arachnoideum( Fig. 6), and sometimes side by side ( e.g., Schwartsburd 3323and 3328; Gutierrez 1133aand 1133b). It clearly resembles young plants of P. esculentumsubsp. arachnoideum ,which have heteroblastic fronds (copiously setosepilose), and yet it is fully fertile with well-developed spores. Ranal (2004)made experiments of germinating spores from tree barks, and the Pteridiumspecimens she grew resulted in such forms: for example Ranal 776,which was cited by Arantes et al.(2008)as “ Pteridium arachnoideum”. We hypothesize that pedomorphy in Pteridiummay be triggered by self-crosses of gametophytes. This morphotype has long been reported in literature, for example by Rosenstock (1907)and Sehnem (1972), as “ Pteridium aquilinumvar. umbrosumChrist ex Rosenst.”, an illegitimate name due to an earlier Pteridium aquilinumvar. umbrosum Luerssen (1889: 107). Rosenstock’s description and typesformed the base for this variety, which we here validate. These specimens are very common in southern Brazil, and similar specimens were also found in Mesoamerica (see Moran 1995: 162). 2. Pteridium esculentum(G.Forst.) Cockayne subsp. campestre(Schrad.) Schwartsb. & J.Prado, comb. nov. et stat.
1920-11
2000-11-24
1920-11
M, L, F, C, LIL, A, NY, J, CAY, P
Manaure & Grant & Cardona & South & Cerro Avila & Vogl & Smith & St. & Massif Dekou-Dekou & de Granville
Colombia
Sierra de Perija
1585
4.7
Kanuku Mountains
1306
-53.95
Mount Auyantepui
7
29
1
Cesar
1932-11
1990-07-16
1932-11
C, A, L, MO, R, S
Locality & Yamapaca & Bues & Yucumo
Bolivia
Ballivian Province
1700
-15.283334
Fay
1285
-67.066666
Fay
7
29
1
La Paz
[271,1380,1744,1769]
1912-10
1912-11-31
1912-10
O
Nord Yungas & Polo-Polo & Coroico
United States of America
1100
Buchtien
7
29
1
1917-12
O
Nord Yungas & Millunguaya
United States of America
1300
Herb. Boliv.
Buchtien
7
29
1
1906-12-03
2006-01-18
1906-12-03
O, M, J
Sud Yungas & Florida
United States of America
Buchtien
2100
-18.239723
Wen
21
-63.67778
Nee
7
29
1
Santa Cruz
1907-10
2008-03-15
1907-10
O, M, L
San Carlos Mapin
United States of America
Buchtien
750
-11.591111
Without
21
-41.2075
Province
7
29
1
Bahia
1944-08-22
1991-09-19
1944-08-22
S, A, H, C, T, R, L, G, M, F, VIES
Santa Cruz de Cabralia & da Contagem
United States of America
Chapada da Contagem
90
-16.383333
Soderstrom
1282
-39.133335
Irwin
7
29
1
Ceara
1988-08-12
O, J, L, VIES
Marechal Floriano
United States of America
700
-20.4131
Behar
7
-40.6831
Pereira
7
29
1
Ceara
2008-09-06
T, S, VIES
United States of America
8
30
-20.133333
Lorencini
1268
-40.816666
Santa Maria de Jetiba
7
29
1
Ceara
1966-02-14
1985-05-17
1966-02-14
W, MBML, H, S
Santa Teresa & Boone
United States of America
Chapada
1000
-14.0
Irwin
41196
-47.0
Veadeiros
8
30
1
Goias
1968-07-04
J, A, UFG
Goiania & Neropolis
United States of America
Rizzo
8
30
1
Goias
1973-05-16
W, R
United States of America
1230
Anderson
8
30
1
1963-09-12
2007-08-18
1963-09-12
P, G, RB, I, A, SP, R, LP, J, Y
Cocalzinho
United States of America
Santos
1150
-25.140833
Prado
21
-50.029446
Ponta Grossa
8
30
1
Mato Grosso do Sul
1949-03-05
2014-07-04
1949-03-05
A, C, L, LP, P, B, VIC, J, G, RB, N, W, T, K, INPA, MG, NY, R, S, SP
Tijucas do Sul & Krapovickas & Cristobal & Boa Vista & Santa Fe
United States of America
-6.424444
Cachoeira de Macacu
21
-35.69917
8
30
1
Rio de Janeiro
1972-12-29
2006-02-23
1972-12-29
G, E, CORD, J, R, Y, SP, A, MO, N, S
Monte Castelo & Lajeadinho & Barboza & Pindamonhangaba
Paraguay
Pico do Itapeva
1888
-25.2
Hirai
1247
-57.116665
Prado
8
30
1
Caaguazu
1941-10-18
2012-02-28
1941-10-18
E, MO, A, LIL, L, RB, BAL
de Azucar & de Mahona & Alo & Mon
Uruguay
Pan de Azucar
194
-31.621973
Sierra La Peregrina
1
-56.042973
Sierra de Mahona
8
30
1
Ciudad Autonoma de Buenos Aires
[802,1447,880,905]
1932-01-29
C, M, SI
Hicken
Uruguay
8
30
1
Ciudad Autonoma de Buenos Aires
1910-05-27
1952-08-26
1910-05-27
A, T, CORD
Castillon & Hunziker
Sierra
El Rodeo
Depto. Calamuchita
8
30
1
[313,1066,988,1013]
SI
Burkart
San Alberto
Sierra
8
30
1
2005-09-03
R, E, CORD
Morero
Depto. San Javier
Los Molles
8
30
1
1889-09-15
1952-06-25
1889-09-15
F, SI, T, M
Corrientes & Petersen
Mburucuya
Sierra de Achala
Kurtz
8
30
1
1936-11-08
1977-01
1936-11-08
A, LP, SI
Dept. General San Martin & Burkart
Tres Cerros
Beron de Astrada
Schinini
8
30
1
1940-03-17
1975-04-27
1940-03-17
A, MO, SI, N, S
Jujuy & Burkart & Troncoso
Termas de Reyes
San Miguel
Schinini
8
30
1
1912-09-19
2002-11-14
1912-09-19
F, O, SI
Zuloaga & Misiones & Dept. Candelaria & Rodriguez
Santa Ana
Depto. Manuel Belgrano
de Zapala
8
30
1
[136,1447,1240,1265]
1991-05-22
Delucchi
Dept. General Manuel Belgrano
Bernardo de Irigoyen
8
30
1
[136,961,1276,1301]
1997-10-15
S, G, SI
Tressens
Dept. Guarani
Predio Guarani
8
30
1
1950-12-16
J, E, LP
Montes
Dept. Iguazu
Puerto Wanda
8
30
1
[509,1324,1312,1337]
1945-07-15
A, LP
Krapovickas
Dep. San Ignacio
Teyucuare
8
30
1
2016-02-21
A, G, LP
Yanez & Marquez
Saint Pierre and Miquelon
Dept.
Ruta Provincial
8
30
1
1922-02-19
2016-02-17
1922-02-19
A, G, LP, L, H, S
Yanez & Marquez & Salta & Camino & Jujuy & Caldera & Babrera & Schwabe & Tucuman & Famailla & La Fronterita & Venturi
United States of America
500
Pinalito Provincial Park
8
30
1
1907-11
1989-04-29
1907-11
O, P, N, A
Buchtien & Alto Caparao & Salino
Bolivia
Charopampa
570
Caparao
Mapini
9
31
1
Minas Gerais
2004-03-01
A, BHCB
Salino
Bolivia
662
-15.805
Mata do Boi Rajado
21
-40.51611
Bandeira
9
31
1
Minas Gerais
1873-09-05
H, BM, K, LE
Mosen & Herb. Bras. Regnell.
Bolivia
Caldas
9
31
1
Minas Gerais
2003-09-13
A, BHCB
Salino
Bolivia
1250
-19.10086
Conceicao do Mato Dentro
1
-43.57414
9
31
1
Minas Gerais
1989-03-13
R, F, CESJ, SPF
Novelino
Bolivia
9
31
1
Minas Gerais
1985-10-05
M, J, T, S, UEC
Chagas
Bolivia
Grandi
Moeda
9
31
1
Minas Gerais
[286,1447,628,653]
2006-09-07
G, BHCB
Heringer
Bolivia
1277
-20.322779
Moeda
21
-43.94167
9
31
1
Minas Gerais
[136,922,664,689]
1998-05-13
P, E, A, BHCB
Ouro Preto & Salino
Bolivia
Itacolomi
9
31
1
Minas Gerais
2004-02-18
RPPN, J, B, C, BHCB
Figueiredo
Bolivia
Rio Acima
Queiroz
Andaime
9
31
1
Minas Gerais
2003-10-10
A, BHCB
Salino
Bolivia
800
-16.404583
Distrito Talisma
1
-40.05761
Santa Maria do Salto
9
31
1
Minas Gerais
2009-06-07
P, B, SP
Sao Thome das Letras & Schwartsburd
Bolivia
Camping do Noel
9
31
1
Minas Gerais
1997-07-12
M, A, HUFU, SP
Ranal
Bolivia
Estacao Ecologica do Panga
Uberlandia
9
31
1
Minas Gerais
[601,1051,808,833]
1999-07-12
M, A, SP
Ranal
Bolivia
9
31
1
Minas Gerais
1999-07-31
M, A, SP
Ranal
Bolivia
9
31
1
Minas Gerais
1999-11-28
2005-04-20
1999-11-28
M, A, SP, P, B
Ranal
Bolivia
Schwartsburd
Bateias
9
31
1
Parana
[322,1164,880,905]
2005-04-24
P, B, MBM
Schwartsburd
Bolivia
Alto da Serra
Bateias
9
31
1
Parana
2007-03-04
P, B, A, M, SP
Schwartsburd & X. Lima
Bolivia
1400
Morro Capivari Grande
9
31
1
Campina Grande do Sul
1988-07-21
M, V, K
Bolivia
Endo
Guaporema
9
31
1
Campina Grande do Sul
[591,1276,952,977]
1988-07-21
M, V, K
Bolivia
Endo
Indianopolis
9
31
1
Campina Grande do Sul
1988-04-23
J, C, S
Bolivia
Pereira
Nova Olimpia
9
31
1
Campina Grande do Sul
[713,1447,988,1013]
1970-03-01
G, MBM
Hatschbach
Bolivia
Pinhais
Piraquara
9
31
1
Campina Grande do Sul
[136,814,1024,1049]
1988-08-17
J
Bolivia
Vituri
Planalto
9
31
1
Campina Grande do Sul
1988-01-27
M, C
Bolivia
Quatro Barras
Antunes
Graciosa
9
31
1
Campina Grande do Sul
[380,1240,1060,1085]
1988-02-25
M, V, K
Bolivia
Rondon
Endo
Indianopolis
9
31
1
Campina Grande do Sul
1988-08-31
C
Alberto Bonezzi
Bolivia
Cerro Negro
Santo Antonio do Sudoeste
9
31
1
Campina Grande do Sul
1988-08-17
M, S
Bolivia
Matzenbacher
Sao Jorge D'Oeste
9
31
1
Campina Grande do Sul
1985-01-26
A, C, L, LP
Krapovickas
Bolivia
Cristobal
9
31
1
Sao Mateus do Sul
1988-01-07
2014-07-04
1988-01-07
M, A, P, B, VIC
Schwartsburd
Bolivia
Sao Miguel
-22.45
Mascarenhas
1260
-42.616665
Dois Vizinhos
9
31
1
Rio de Janeiro
1965-11-04
P, N, R, M, A, F
Itatiaia & Tryon
Bolivia
Itatiaia
Tryon
Abrigo Reboucas
9
31
1
Rio de Janeiro
1911-10
B, L, C, A, S, H
Luederwaldt
Bolivia
Locality
Santa Cruz
Anonymous
9
31
1
Rio Grande do Sul
syntype
1907-08
1911-10
1907-08
H
Luederwaldt
Bolivia
Hammonia
Sao Paulo
Luederwaldt
9
31
1
Sao Paulo
syntype
1910-06
H
Luederwaldt
Bolivia
Municipality
Estacao Alto da Serra
Santo Andre
9
31
1
Sao Paulo
syntype
W, J, B, K, LE
United States of America
Municipality
Rosenstock Filic.
Anonymous
9
31
1
syntype
W, J, B, K
Locality
United States of America
9
31
1
syntype
1948-09-11
1988-06-23
1948-09-11
A, BM, MBM, E, MO
Macedo & Schinini & Zardini
Paraguay
-26.05
Parque Nacional Ybycui
1243
-57.0
Locality
9
31
1
Caaguanzu
syntype
1927-11
1988-04-06
1927-11
S, LP, SI, C, M, MO
Meyer
Uruguay
Dep. Misiones
Mar del Plata
Isla San Pablo
9
31
1
Ciudad Autonoma de Buenos Aires
syntype
1928-03
2016-11-29
1928-03
T, SI, BA
Meyer & Gutierrez
Uruguay
Fontana
Villa Benegas
Camino
9
31
1
Cordoba
syntype
T, M, LP
Los Molles & Castellano & Dept. Mburucuya & Estancia Santa Teresa
Uruguay
Petersen
9
31
1
Corrientes
syntype
1949-05-16
1979-09-16
1949-05-16
A, LP, J, E
Schinini & Montes
Uruguay
Dept. San Martin
Cerro de Susini
Tres Cerros
9
31
1
Misiones
syntype
1951-05-22
A, L, LP
Cabrera
Uruguay
Mayo
Colonia
9
31
1
Misiones
syntype
[249,814,1888,1913]
1918-07-20
G, T, SI
Bertoni
Uruguay
Dept. Iguazu
9
31
1
Misiones
syntype
1901-10-21
T, CORD
Stuckert
Uruguay
Dept. Montecarlo
Alto Parana
Puerto Piray
9
31
1
Misiones
syntype
[522,1316,1924,1949]
1937-07-11
BA
Perez Moreau
Uruguay
Teyu Cuare
Dept. San Ignacio
9
31
1
Misiones
syntype
1922-11
A, G, LP, S, BA
San Pedro & Esmeralda Provincial Park & Yanez & Famailla & La Fronterita & Venturi
Uruguay
Marquez
9
31
1
Tucuman
syntype
[457,1025,1996,2021]
1911-12
L, SI
Castillon
Uruguay
Valle de Tafi
9
31
1
Tucuman
syntype