Pteridium arachnoideum (Kaulf.) Maxon subsp. arachnoideum

Schwartsburd, Pedro B., De Moraes, Pedro L. R. & Lopes-Mattos, Karina L. B., 2014, Recognition of two morpho-types in eastern South American brackens (Pteridium-Dennstaedtiaceae-Polypodiopsida), Phytotaxa 170 (2), pp. 103-117: 109-112

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http://doi.org/ 10.11646/phytotaxa.170.2.3

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http://treatment.plazi.org/id/F554879F-FFFE-A656-E8BB-BF10FBA8FBB5

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Felipe

scientific name

Pteridium arachnoideum (Kaulf.) Maxon subsp. arachnoideum
status

 

1. Pteridium arachnoideum (Kaulf.) Maxon subsp. arachnoideum   . ( Figs. 1D–F View FIGURE 1 , 2A–C View FIGURE 2 )

Pteridium arachnoideum (Kaulf.) Maxon (1924: 89)   . Pteris arachnoidea Kaulfuss (1824: 190)   , nom. cons. Allosorus arachnoideus (Kaulf.) Presl (1836: 153)   , as “(Desv.) C. Presl”. Pteris aquilina Linnaeus (1753: 1705) var. arachnoidea (Kaulf.) Eaton (1861: 203)   . Cincinalis arachnoidea (Kaulf.) Trevisan de Saint-Léon (1874:31)   . Pteridium aquilinum   (L.) Kuhn (1879:11) var. esculentum ( Forster 1786: 74) Kuhn (1882: 347) f. arachnoideum (Kaulf.) Hieronymus (1909: 246)   . Pteridium aquilinum   (L.) Kuhn var. arachnoideum (Kaulf.) Brade (1920: 56)   . Filix-foemina aquilina   (L.) Farwell (1931: 290) var. arachnoidea (Kaulf.) Farwell (1931: 290)   . Pteridium caudatum ( Linnaeus 1753: 1705) Maxon (1901: 631) subsp. arachnoideum (Kaulf.) Lellinger (2003: 150)   . Pteridium esculentum (G. Forst.) Cockayne (1908: 34) subsp. arachnoideum (Kaulf.) Thomson (2012: 45)   . Lectotype (designated here):— BRAZIL. “Habitat in Brasília”, without date, C.A. de Chamisso s.n. (LE!). ( Figs. 1D–F View FIGURE 1 , 2A–C View FIGURE 2 )

= 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 Janeiro Brasilia 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!, isolectotype W!).

= Pteris psittacina Presl (1822: 185)   , nom. rej. Allosorus psittacinus (C. Presl) Presl (1836: 153)   , as “ psitaccinus ”. Pteris aquilina   L. 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 Janeiro Brasilia” [Circa Rio de Janeiro in collibus], [December 1818], J.B.E. Pohl s.n. (lectotype PRC!, isolectotype W!).

= Cheilanthes aquilinaris Fée (1873: 37   , t. 91, fig. 1). Hypolepis aquilinaris (Fée) Christ (1901: 636)   . Lectotype (designated here):— BRAZIL. Rio de Janeiro: “Brasilia fluminensis”, [Haut Itatiaia], [6 June 1871], A.F.A. Glaziou 5330 (lectotype B!-20 0074529, isolectotypes C- 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!]). syn. nov.

= Pteridium aquilinum   (L.) Kuhn var. umbrosum H. Christ ex Rosenstock (1907: 90)   , nom. illeg. (non Luerssen 1889: 107). Syntypes:— BRAZIL. Rio Grande do Sul: Santa Cruz, Fazenda Tangerina, withou date, L.C. Jürgens & A. Stier 95 (S? n.v.), Mun. Rio Pardo, Fazenda Soledade, without date, L.C. Jürgens & A. Stier 118 (NY [image!], S? n.v., possible syntype MVM! [“ Jürgens & Stier 190 b ”]). syn. nov.

Plants terrestrial, thicket forming. Rhizomes long-creeping, branching, with two circles of perforated dictyosteles, lanose. Fronds (1.2–)2–3(–4.5) × (1–)1.3–1.8(–3) m, petioles stramineous, proximally with epipetiolar roots, adaxially sulcate, up to 2 m long, glabrous, laminae rhombic, proximally 3–4-pinnate-pinnatifid, medially less dissected, distally pinnatifid, pinnae and pinnules distally with free lobes between the segments, compound distal segments inequilateral, irregularly dissected, caudate at apex, simple distal segments linear, up to 3(–4) cm long, costae abaxially terete, with reddish catenate hairs, 5–7-celled, 0.3–0.5 mm long, adaxially glabrous, costules abaxially strongly flattened with wing-like appearance, with reddish catenate hairs, adaxially glabrous, veins abaxially strongly raised, lanose, with lax arachnoid hairs, 0.6–0.8 mm long, adaxially glabrous, laminar tissue between the veins abaxially glabrous and totally visible, adaxially glabrous, stomatal guard-cells 27–34 µm long, infertile pseudo-indusia 0.2–0.3 mm wide, the marginal cells horizontal.

Distribution and ecology: —Eastern Brazil, Uruguay, Paraguay and central/north-eastern Argentina. From sea level to 2200 m elev., in open habitats (Pampas, Campos Gerais, altitudinal fields, coastal sandy Restinga, crop lands, pastures, disturbed areas), forming extensive thickets, rarely in the edge of forested areas and clearings ( Fig. 3 View FIGURE 3 ).

Representative specimens examined: — BRAZIL. Ceará: Pacoti, Serra do Baturité , [-4.2832°, -38.9314°], 700- 800 m, 9 April 2011, P. B   . Schwartsburd & M. F   . Moro 2490 ( NY, SP, VIC)   . Paraíba: Alhandra , [-7.4397°, -34.9157°], 16 December 1985, A. J. R   . Silva s.n. ( UFP 7135 View Materials )   . Pernambuco: Jaqueira, Usina Colônia, Córrego do Guariba e Serra do Quengo , 08°43’S, 35°50’20”W, 700 m, 18 March 2011, P. B GoogleMaps   . Schwartsburd & Seu Dejau 2342 ( SP, UFP, VIC)   . Alagoas: Ibateguara, Engenho Coimbra , [-9.0114°, -35.8866°], 380–400 m, 19 December 2000, M. R   . Pietrobom-Silva & A. C. P   . Santiago   4680 (UFP). Bahia: Near Poções, [Lucaia], [11] March 1817, Wied s.n. (BR-5418671, K, MEL-2353758). Distrito Federal : 20 km East of Brasília, Chapada da Contagem, [-15.8732°, -48.1567°], 700–1000 m, 19 August 1964, H. S   . Irwin & T. R   . Soderstrom 5301 ( NY, SP)   . Minas Gerais: Alto Caparaó , P. N   . Caparaó , [-20.4080°, -41.8344°], 1970 m, 11 July 2009, P. B   . Schwartsburd et al. 2092 ( VIC)   . Mato Grosso do Sul: Ivinhema , [-22.2360°, -53.7825°], 22 November 2003, A   . Sciamareli et al. 06 ( UPCB)   . Rio de Janeiro: Itatiaia , P. N   . Itatiaia , 22°21’46”S, 44°43’40”W, 2200 m, 14 July 2009, P. B GoogleMaps   . Schwartsburd et al. 2128 ( NY, VIC)   . São Paulo: Ilha dos Alcatrazes , [-24.1005°, -45.6952°], 50 m, 26 October 2011, P. B   . Schwartsburd & L   . Rossi 2570 ( SP, VIC)   . Paraná: Ponta Grossa , P. E   . Vila Velha, Fortaleza , [-25.2305°, -49.9927°], 5 March 2006, P. B   . Schwartsburd et al. 1008 ( UPCB)   . Santa Catarina: Praia Grande , P. N   . Aparados da Serra, [-29.2025°, -50.0321°], 200 m, 5 August 2009, P. B   . Schwartsburd et al. 2255 ( NY, SP, VIC)   . Rio Grande do Sul: São Leopoldo , [-29.7640, -51.1825 °], 22 January 1933, A GoogleMaps   . Sehnem s.n. (SP-50626).— URUGUAY. Artigas: Cuaró , [-30.6174°, -56.9125°], December 1924, C   . Osten 17349 ( MVM)   . Durazno: En   Rio Negro, [-33.0460°, -57.0656°], 3 August 1922, C   . Osten 16517 ( MVM)   . Treinta y Tres: Yerbal , [-33. 2244°, -54.4026°], November 1819, M. B   . Berro 1263 ( MVM)   . Lavalleja: Minas, Arequita , [-34.2865°, -55.2642°], 7 November 1819, M. B   . Berro 1264 ( MVM)   . Flores: Cueva del Tigre, en Guaycurú , [-33.4862°, -56.8385°], 30 October 1896, C   . Osten 3297 ( MVM)   . San José: Serra Mahoma, [-34.0898°, -56.9454°], 1939, Chebataroff 2607 ( MVM)   . Maldonado: Cerro del Pan de Azucar , [-34.8100°, -55.2590°], 28 March 1885, Anonymous [Arechavaleta?] (MVM-4198)   .— PARAGUAY. Amambay: National Park Cerro Corá , 22°39’51”S, 55°19’18”W, 350 m, without date, E. M GoogleMaps   . Zardini & P   . Báez 52308 ( NY)   . Caazapá: Estancia Tapytá of Shell Forestry Ltd. , 26°16’26”S, 55°45’47”W, 150 m, 14 December 1999, E. M GoogleMaps   . Zardini & P   . Báez 52891 ( NY)   .— ARGENTINA. Misiones: Aristóbulo del Vale, Reserva UNLP   , 8 January 2003, G   . Marquez 25 ( VIC). Córdoba: Sierra Grande , [-31.4857°, -64.7422°], 1 February 1936, A. P   . Rodrigo 594 ( NY)   .

Remarks:— Plants from western South America (including north-western Brazil) and Central America, commonly ascribed to Pteridium arachnoideum   s.l., are morphologically different and shall be treated in at least one other subspecies of P. arachnoideum   . These plants present the laminar tissue between the veins abaxially with farinose appearance, not visible, copiously furnished with gnarled hairs, whereas in all 191 specimens of P. arachnoideum subsp. arachnoideum   s. str. analysed, the gnarled hairs are totally absent. Instead, this part is visible and completely glabrous ( Figs. 1F View FIGURE 1 , 2B View FIGURE 2 ), conflicting with the descriptions of Moran (1995), Thomson & Martin (1996: Figs. 4B, 5E, F), and Mickel & Smith (2004). Although Thomson & Martin (1996) regarded P. aquilinum var. arachnoideum   in a broad sense, they indeed observed the same pattern (on page 46): “In contrast, accessions of var. arachnoideum   from three localities in Brazil (...) and one from Uruguay (...) lacked intervein hairs of any kind on the lower surface...”.

In central South America, there is a dry region named the Dry Diagonal (or Diagonal of Open Areas) connecting the Argentinian/Bolivian Chaco, through the Cerrado (Brazilian savannah), up to the Caatinga (Brazilian semi-desert vegetation), and separating South America into two blocks of wetter regions (mainly dominated by forested areas) ( Morrone & Coscarón 1996, Fig 3 View FIGURE 3 ). Pteridium arachnoideum   s.l. is practically absent in the Dry Diagonal, even though it is a species from open areas (personal observation throughout central Brazil, see also Tryon 1941: Map 12), probably requiring wetter environmental conditions to stablish populations. Apparently, P. arachnoideum subsp. arachnoideum   s. str. and the subspecies from western South America and Central America are geographycally segregated by the Dry Diagonal.

2. Pteridium arachnoideum (Kaulf.) Maxon subsp. campestre (Schrad.) Schwartsb. & P.L.R. Moraes   , comb. et stat. nov. ( Figs. 1A–C View FIGURE 1 , 2D–G View FIGURE 2 )

Pteris campestris Schrader (1824: 871)   . Lectotype (designated here):— BRAZIL. Bahia: “Brasilia, Prov. Bahiensis”, [near Ilhéus], 1817, Wied s.n. (lectotype LE! [ex Herb. Schrader], isolectotypes BR!-5419005 [ex Herb. Martii], BR!-5418022 [fragment,ex Herb. l’État]). ( Figs. 1A–C View FIGURE 1 , 2D–G View FIGURE 2 )

= Aquilina gardneri C. Presl ex Ettingshausen (1865: 91   , t. 53, fig. 6, t. 54, fig. 3). Pteris gardneri C. Presl ex Ettingshausen (1864: 42)   , nom. illeg., non Pteris gardneri ( Fée 1852: 139) Hooker (1858   [1852]: 206). Type:— BRAZIL. State unknown: “In Brasilia”, without date, G. Gardner, coll. Pl. Bras. 1224 (holotype W- n.v., isotype PRC! [ex Herb. Presl]). syn. nov.

Plants terrestrial, thicket forming. Rhizomes long-creeping, branching, with two circles of perforated dictyosteles, lanose. Fronds 1.0–1.5(–2.0) × 0.6–1.0 m, petioles stramineous, proximally with epipetiolar roots, adaxially sulcate, up to 0.8 m long, glabrous, laminae rhombic, proximally 3-pinnate-pinnatifid, medially less dissected, distally pinnatifid, pinnae and pinnules distally without free lobes between the segments, compound distal segments equilateral, regularly dissected, not caudate at apex, simple distal segments linear, up to 1.0(–1.5) cm long, costae abaxially terete, with reddish catenate hairs, 5–7-celled, 0.3–0.5 mm long, adaxially glabrous, costules abaxially slightly flattened, with reddish catenate hairs, adaxially glabrous, veins abaxially slightly raised, hirsute, with conspicuous stiff acicular hairs, 0.2–0.3 mm long, adaxially glabrous, laminar tissue between the veins abaxially with farinose appearance, not visible, totally covered by conspicuous gnarled hairs, adaxially glabrous, stomatal guard-cells 28–35 µm long, infertile pseudoindusia 0.3–0.4 mm wide, the marginal cells vertical.

Distribution and ecology: —Endemic to north-eastern Brazil, at or near sea level in open habitats with white sand soil (Restinga and Mussununga; Saporetti-Junior et al. 2012), a few collections inland up to ca. 1000 m, also from sandy soil spots ( Fig. 3 View FIGURE 3 ).

Representative specimens examined: — BRAZIL. Ceará: Queimadas, São Benedito, Serra da Ibiapaba , [- 7.1021°, -40.1778°], 7 January 1942, P   . Bezerra 403 ( IPA)   . Paraíba: Mamanguape , BR 101   , próximo ao km 55, [- 6.8526°, -35.1327°], 17 September 1979, P   . Martins et al. s.n. ( EAC 6936 View Materials , UFP 7218 View Materials )   . Pernambuco: Igarassu, Refúgio Ecológico Charles Darwin , 07°48’37”- 07°49’27”S, 34°27’25”- 34°56’52”W, 18 October 1999, A. C. P GoogleMaps   . Santiago   38 (PEUFR). Bahia: Jacobina, Monte Taboa , ca. 2 km da cidade, 11°11’26”S, 40°30’33”W, 656 m, 3 August 2001, F. R GoogleMaps   . Nonato et al. 927 ( HUEFS, SP, VIC)   .

Remarks: —Apart from us, some authors from the 19 th Century had recognized this taxon as distinct from the other South American brackens, for example: Schrader (1824 —as Pteris campestris   ), Presl (herbarium identifications—as Aquilina gardneri   ), and Ettingshausen (1864, 1865 —as Pteris gardneri   and A. gardneri   ).

Pteridium arachnoideum subsp. campestre   is a smaller taxon than P. arachnoideum subsp. arachnoideum   s. str., forming much smaller thickets. It has smaller fronds (1–1.5 × 0.6–1 m vs. 2–3 × 1.3–1.8 m), 3-pinnate-pinnatifid laminae (vs. 3–4-pinnate-pinnatifid), pinnae and pinnules without free lobes between the distal segments (vs. free lobes always present), compound distal segments equilateral, regularly dissected, not caudate at apex (vs. inequilateral, irregularly dissected and caudate at apex), simple distal segments up to 1(–1.5) cm long (vs. to 3(–4) cm long), costules abaxially slightly flattened (vs. strongly flattened, with wing-like appearance), veins abaxially slightly raised and hirsute, with stiff acicular hairs, 0.2–0.3 mm long (vs. strongly raised and lanose, with lax arachnoid hairs, 0.6–0.8 mm long), laminar tissue between the veins abaxially with farinose appearance, not visible, copiously furnished with gnarled hairs (vs. without farinose appearance, visible and glabrous), infertile pseudo-indusia 0.3–0.4 mm wide with the marginal cells vertical (vs. 0.2–0.3 mm wide, marginal cells horizontal) ( Figs. 1A–C View FIGURE 1 , 2E–G View FIGURE 2 vs. Figs. 1D–F View FIGURE 1 , 2B, C View FIGURE 2 , and see also Ettingshausen 1865: t. 53, 54).

Pteridium caudatum   also occurs in Brazil but is restricted to the Amazonian region in the north-west ( Tryon 1941, Prado & Moran 2009). Besides geography and the distinguishing characteristics given in the key, P. arachnoideum subsp. campestre   differs from P. caudatum   by compound distal segments equilateral, regularly dissected and not caudate at apex (vs. inequilateral, irregularly dissected and caudate), and by simple distal segments which are up to 1(–1.5) cm long (vs. to 4(–7) cm long) ( Figs. 1A–C View FIGURE 1 , 2E–G View FIGURE 2 ).

Pteridium arachnoideum subsp. campestre   differs from P. aquilinum subsp. pseudocaudatum ( Clute 1900: 39) Hultén (1941: 44)   by compound distal segments equilateral, regularly dissected and not caudate at apex (vs. inequilateral, irregularly dissected and caudate), simple distal segments that are up to 1(–1.5) cm long (vs. to 3 cm), and laminar tissue between the veins that is abaxially furnished with gnarled hairs (vs. glabrous). Finally, Pteridium arachnoideum subsp. campestre   differs from P. aquilinum subsp. feei (W. Schaffn. ex Fée 1857: 73) Thomson et al. (2008: 13)   by laminar tissue between the veins abaxially furnished with gnarled hairs (vs. glabrous) and infertile pseudo-indusia with glabrous margins (vs. margins ciliate) ( Figs. 1A–C View FIGURE 1 , 2E–G View FIGURE 2 ).

P

Museum National d' Histoire Naturelle, Paris (MNHN) - Vascular Plants

B

Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet

M

Botanische Staatssammlung München

F

Field Museum of Natural History, Botany Department

NY

William and Lynda Steere Herbarium of the New York Botanical Garden

SP

Instituto de Botânica

VIC

Universidade Federal de Viçosa

A

Harvard University - Arnold Arboretum

J

University of the Witwatersrand

R

Departamento de Geologia, Universidad de Chile

UFP

Universidade Federal de Pernambuco

C

University of Copenhagen

K

Royal Botanic Gardens

H

University of Helsinki

S

Department of Botany, Swedish Museum of Natural History

T

Tavera, Department of Geology and Geophysics

N

Nanjing University

UPCB

Universidade Federal do Paraná

L

Nationaal Herbarium Nederland, Leiden University branch

E

Royal Botanic Garden Edinburgh

G

Conservatoire et Jardin botaniques de la Ville de Genève

IPA

Empresa Pernambucana de Pesquisa Agropecuária, IPA

HUEFS

Universidade Estadual de Feira de Santana

Kingdom

Plantae

Phylum

Tracheophyta

Class

Polypodiopsida

Order

Polypodiales

Family

Dennstaedtiaceae

Genus

Pteridium

Loc

Pteridium arachnoideum (Kaulf.) Maxon subsp. arachnoideum

Schwartsburd, Pedro B., De Moraes, Pedro L. R. & Lopes-Mattos, Karina L. B. 2014
2014
Loc

Pteridium arachnoideum (Kaulf.)

Thomson, J. A. 2012: )
Farwell, O. A. 1931: 290
Farwell, O. A. 1931: )
Maxon, W. R. 1924: )
Brade, A. C. 1920: )
Maxon, W. R. 1901: 1705
Kuhn, M. 1879: 11
Presl, C. B. 1836: )
Kaulfuss, G. F. 1824: )
Forster, J. G. A. 1786: 74
Eaton, D. C. 1753: )
1924
Loc

Cheilanthes aquilinaris Fée (1873: 37

Christ, H. 1901: )
Fee, A. L. A. 1873: 37
1873
Loc

Pteris campestris

Schrader, H. A. 1824: )
1824
Loc

Aspidium brasilianum

Kuhn, M. 1882: )
Presl, C. B. 1836: )
Presl, C. B. 1822: )
1822
Loc

Pteris psittacina

Christensen, C. 1934: )
Maxon, W. R. 1933: )
Baker, J. G. 1870: )
Presl, C. B. 1836: )
Presl, C. B. 1822: )
1822