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Actu Biol vol.35 no.98 Medellín ene./jun. 2013
ART&ICULOS DE INVESTIGACI&ON
Novelties in Grammitis Sw. and StenoGrammitis Labiak (Polypodiaceae) from Costa Rica, Colombia, and Venezuela
Novedades en Grammitis Sw. Y StenoGrammitis Labiak (Polyp odiaceae) para Costa Rica, Colombia y Venezuela
Alexander F. Rojas-Alvarado1
Escuela de Ciencias Biol&gicas. Universidad Nacional de Costa Rica. Apartado postal: 86-3000. Heredia, Costa Rica. Correo electr&nico: .
Recibido: enero de 2012; aceptado: noviembre de 2012.
Four new species of Grammitis Sw. (G. angustissima A. Rojas, G. roraimensis A. Rojas, G. spathulata A. Rojas
and G. undulata A. Rojas) and one new species of StenoGrammitis Labiak (S. grammitoides A. Rojas) are described
here. The first species is known only from the Colombian Andes and is characterized by the
second is known only from the Roraima Mountains in Venezuela, and is the largest species with a long ascending
rhizome and a branched b the third species is similar to G. marginella (Sw.) Sw. in spathulate
blade with rounded apex, but has sessile fronds
the fourth species is also known from the
Colombian Andes, and it is similar to G. angustissima in its narrow blade, but has undulate margins and a narrower
blac the StenoGrammitis species is endemic to Cocos Island, Costa Rica, and is characterized by clathrate rhizome scales, an entire blade, and discontinuous sori that are oblique at the costa.
Key words: Grammitis, grammitoids, Neotropic, new taxa, Polypodiaceae, StenoGrammitis.
Cuatro especies nuevas de Grammitis Sw. (G. angustissima A. Rojas, G. roraimensis A. Rojas, G. sphatulata
A. Rojas and G. undulata A. Rojas) y una nueva especie de StenoGrammitis Labiak (S. grammitoides A. Rojas)
son descritas aqu&. La primera especie es conocida solo para los Andes colombianos y se caracteriza por l& la segunda especie es conocida s&lo para las monta&as de Roraima en Venezuela, y es la especie
m&s grande con rizoma largo ascendente y cinta negra del margen de la l& la tercera especie es
similar a G. marginella (Sw.) Sw. en l&mina espatulada con &pice redondeado, pero tiene frondas s&siles y l& la cuarta especie es tambi&n conocida para los Andes colombianos y es similar a G. angustissima en la
l&mina delgada, pero tiene margen ondulado y cinta negra del margen de la l&mina m& la especie de
StenoGrammitis es end&mica de la Isla del Coco, Costa Rica, y es caracterizada por escamas del rizoma clatradas, l&mina entera y soro discontinuo y oblicuo a la costa.
Palabras clave: Grammitis, grammitoides, nuevos taxones, neotr&pico, Polypodiaceae, StenoGrammitis.
INTRODUCTION
The genus Grammitis Sw. includes those
species with simple and entire fronds, with
a marginal band of black or dark brown
sclerenchyma (Moran and Smith 1995). Moran
and Smith (1995) mentioned that although the
genus (Grammitis) has been revised tree times
in this century (Bishop 1977, Copeland 1952,
Maxon 1915), few specimens were available
from the studies. Many recent collections,
some of which have been described as new
species, make necessary a new revision of the genus.
Labiak (2011) mentioned that StenoGrammitis
Labiak is readily distinguished from other
grammitid genera by its linear laminae, often not
reaching more than 0.5 cm in width, clathrate
and iridescent rhizome scales, segments with
a single and unbranched vein, and by having a single sorus per segment.
The genus StenoGrammitis differs from the
related genus Lellingeria A.R. Sm. & R.C.
Moran by linear leaves usually less than 5 mm
wide, clathrate iridescent rhizome scales
that are glabrous except for a single apical
cilium, veins unbranched and only one per
segment, fertile veins usually with the dark
sclerenchyma visible beneath the sporangia, and x = 33 (Labiak 2011).
Based on the author's studies of the Neotropical
ferns and revised specimens in COL, CR,
MO, US y USJ herbaria, five new species are described here.
MATERIALS AND METHODS
The deposit of the respective types and
specimens examined of each taxon treated
here is specified in the section of each taxon.
The abbreviations used here (herbarium
code) correspond to the following herbaria:
COL = Herbario Nacional Colombiano,
Universidad Nacional de Colombia (Bogot&
D. C., Colombia); CR = Herbario Nacional,
Museo Nacional (San Jos&, Costa Rica);
MO = Missouri Botanical Garden (Saint
Louis, Missouri, U. S. A.); NY = New York
Botanical Garden (Bronx, New York, U. S. A.);
US = United States National Herbarium,
Smithsonian Institution (Washington D. C.,
U. S. A.); USJ = Herbario ''Luis A. Fournier'',
Universidad de Costa Rica (San Pedro, San
Jos&, Costa Rica).
RESULTS AND DISCUSSION
New species
Grammitis angustissima A. Rojas, sp. nov.
TYPE: COLOMBIA: Putumayo: 50 km W de
El Pepino, 2,050 m, 20 Nov 1972, W. Hageman and Leist 1496 (Holotype: COL).
Novum taxon Grammiti bryophila simile, sed
lamina foliari angustiora, crassiora, margine
nigrescenti, latiore, venis inconspicuis in sterili parte dignocendum.
E rhizome 0.5-1 mm in diameter, rhizome scales 2.5-3.5
x 0.3-0.5 mm, linear-lanceolate, brown, 6-10 cells
in broad at base, fronds
2.5-8.7 cm long, pendulous, stipe 4-7 mm
long, dark brown, blade 2.1-8 x 0.15-
0.25 cm, linear, chartaceous, sparsely hairy
adaxially, glabrous at margin, the hairs ca. 0.2
mm long, 2-4-celular, uniseriate, with a dark
elonged apical cell, blackish marginal ribbon
0.1-0.2 main vein dark brown to
atropurpureous abaxially, green to light brown
adaxially or darker only at base, promi sterile veins single, few evident
to inconspicuous, fertile veins very evident,
oblique at 25-40& w sori
1-1.5 mm long, rounded or slightly elongated,
sometimes confluent at maturity, present in the 2/5 distal portion of the blade.
Distribution. Known only from the type collection.
Etymology. The specific epithet refers to its narrow blade.
Diagnosis. Grammitis angustissima differs from
G. bryophila (Maxon) F. Seym. by its narrower
(1.5-2.5 mm broad vs. (3-) 4-7 mm) or (14-)
20-32 times longer than broad (vs. 7-17 times)
and thicker (chartaceous vs. papyraceous to
subchartaceous) blade with broader (0.1-0.2 mm
broad vs. ca. 0.1 mm) blackish marginal ribbon,
and not evident (vs. evident) veins in sterile portion ().
Grammitis roraimensis A. Rojas, sp. nov.
TYPE: VENEZUELA: Bolivar: summit of
mount Roraima, on NW portion, N and NW of Summit camp, 2,620-2,740 m, 27 Sep 1944, J. Steyermark 58 837 (Holotype: US).
Nova species Grammitidi paramicola similis,
sed frondibus magnioribus et rhizomatibus longi-ascendentibus separatur.
T rhizome ca. 1 mm in diam., long
ascending to 4.5 rhizome scales
0.5-1.5 x 0.2-0.3 mm, lanceolate, 5-8 cells in
broad at the base, fronds 18-
27 cm long, erect, stipe 10-15 mm long,
atropurpureous, winged and continuous with
the blade, blade 17-29 x 0.3-0.6 cm,
linear-sphatulate to linear, chartaceous, acute
at apex, glabrous, blackish marginal ribbon
0.2-0.3 mm broad,
vein light green to brown, few evident abaxially,
v sterile veins single,
inconspicuous, fertile veins single or bifurcate,
few evident, oblique at 15-40& wit sori 0.5-1.5 mm broad, oblong, confluent
at maturity, present only in the 1/5 distal portion of the blade.
Distribution. Known only from the type collection.
Etymology. The specific epithet refers to its geographic distribution.
Diagnosis. Grammitis roraimensis differs
from G. paramicola L. E. Bishop by its longer
(20-45 mm long vs. 3-10 mm) rhizome, longer
(17-29 cm long vs. 2-7.5 cm) blade and branched
(vs. no branched) blackish marginal ribbon ().
Grammitis spathulata A. Rojas, sp. nov.
TYPE: COLOMBIA: Magdalena: Sierra de
Perij&, Casacar& Valley, 23 km E of Codazzi,
2 km from the Venezuelan border, 2,550 m, 12
Feb 1945, M. Grant 10 943 (Holotype: COL; Isotype: US).
Species haec nova stipem non exhibet, lamina
foliaris sua glabra est. Per hos characteres
nova species a Grammitidi marginella distinguenda est.
E rhizome 0,6-0,9 mm in diam.,
compact, rhizome scales 0.5-1.5 x
ca. 0.3 mm, linear-lanceolate, 6-10 cells broad
at the base, fronds 2.2-5.5 cm long, blade 0.4-0.8 cm broad,
oblanceolate to spathulate, chartaceous,
rounded at apex, glabrous in both surfaces,
blackish marginal ribbon 0.2-0.3
main vein dark brown to atropurpureous
basally, light brown distally, evident abaxially,
f sterile and fertile veins
single to bifurcate, few evident, oblique at
10-25& w sori 2.5-3.5 mm
long, rounded to ovate, few times confluent
at maturity, present in the 1/2 distal portion of the blade.
Distribution. Known only from the type collection.
Etymology. The specific epithet refers to its spathulate blade.
Diagnosis. Grammitis spathulata differs from
G. marginella (Sw.) Sw. in its absent (vs.
present) stipe and glabrous (vs. hairy) blade
(). For the sessile fronds no other species are similar.
Grammitis undulata A. Rojas, sp. nov.
TYPE: COLOMBIA: Nari&o-Junin: 5 km en
direcci&n de Barbacoas, 1,000 m, 20 Dec 1972, W. Hagemann and Leist 1759 (Holotype: COL).
Nova species haec Grammitidi angustissimae
affinis est, sed prior rhizomatis squamis
brevioribus, lamina foliari tenuiora, in margine
undulata, vitta marginali atropurpurea,
angustiora, nervis patentibus in sterili parte foliorum distinguenda est.
E rhizome 0.5-0.7 mm in diam.,
compact, rhizome scales 0.5-1 x 0.1-
0.2 mm, linear-lanceolate, 4-7 cells in broad at
the base, fronds (4-) 6.5-13 cm
long, pendulous, stipe (1-) 4-15 mm
long, brown, blade (4-) 6-11.5 x 0.15-
0.25 cm, linear, papyraceous to subchartaceous,
glabrous in both surfaces, blackish marginal
ribbon 0.5-0.1 main vein dark brown
to atropurpureous basally, light brown distally,
evident abaxially, inconspicuous adaxially, not
pro sterile and fertile veins
single to bifurcate, few evident, oblique at 5-15&
w sori 1-2 mm long, rounded
to ovate, sometimes confluent at maturity, present in the 1/2 distal portion of the blade.
Distribution. Colombia, Andean Cordillera, at 600-1,000 m alt.
Etymology. The specific epithet refers to its undulate blade.
Paratype. COLOMBIA. Nari&o: carretera
Junin-Barbacoas, hacia Buena Vista, 600 m, 20 Dec 1972, W. Hagemann and Leist 1760 (COL).
Diagnosis. Grammitis undulata differs from
G. angustissima A. Rojas by its smaller (0.5-
1(-1.5) x 0.1-0.2 mm vs. 2.5-3.5 x 0.3-0.5 mm)
rhizome scales, undulate (vs. entire) and thinner
(papyraceous vs. chartaceous) blade with narrower
(0.05-0.1 mm broad vs. 0.1-0.2 mm) blackish
marginal ribbon, main vein not prominent (vs.
prominent) and secondary veins evident (vs. no evident) in sterile portion ().
StenoGrammitis grammitoides A. Rojas, sp. nov.
TYPE: COSTA RICA: Puntarenas:
Puntarenas, Parque Nacional Isla del Coco,
orillas del R&o Genio, Los Llanos, en el sendero
a Cerro Iglesias, 5&32'05''N, 87&03'40''W,
240-260 m, 6 Jan 2010, A. Rojas y J. Chaves
8946 (Holotype: CR; Isotypes: MO, USJ).
Nova species Cochlidio proctorii propinqua,
sed laminae foliaris margine brunnea (ad vicem
virescenti), sine pilis (ad vicem pilosa), soris
mediis (ad vicem supracostalibus) et angulo
25&-40& (ad vicem 5&-15&) ad costam respectu
inclinatis distinguenda.
E rhizome 0.4-0.7 mm in diam., rhizome scales 0.5-1 x 0.4-0.6 mm,
ovate, clathrate, 5-8 cells in broad at the base,
fronds 4-11.5 cm long, stipe
absent or less than 2.5 mm long, atropurpureous,
hairy, the hairs ca. 0.2 mm long, brown-gold,
s blade 4-11.5 x 0.3-0.5 cm,
linear-elliptic to rarely linear, glabrous, without
blackish margin or it less than 0.05 mm and
brown, main vein dark brown to
blackish, hairy, the hairs 0.1-0.2 mm, single to
fasciculate, sometimes bifurcate with a branch
long sterile and fertile veins
single, inconspicuous, oblique at 25-40& wit hydathodes 1-2 x 0.2-0.4 mm,
oblong to elliptic,
sori ca. 0.5
mm broad, 1-2 mm long, oblong, no confluent
at maturity, present in the 1/3 distal portion of
the blade.
Distribution. Costa Rica, known only from the
type from Cocos Island.
Etymology. The specific epithet makes reference
to its simple bade with oblique sori and
sometimes a dark marginal blade ribbon same
as in the genus Grammitis.
Diagnosis. StenoGrammitis grammitoides differs
from other species of the genus because have
entire blade, but is included here because
have clathrate rhizome scales, inequilaterally
branched costal hairs and hydathodes present.
The new species differs from the Jamaican
Cochlidium proctorii (Copel.) L.E. Bishop by
its brown (vs. green) and entire (vs. repand)
blade margins, without (vs. with) marginal hairs,
medial (vs. supracostal) sori and they oblique
at 25-40& (vs. 5-15&) with respect to costa
(figure 1, M-O). Due to the relation of the new
species with Cochlidium proctorii, probably the
second need to be combined under the genus
StenoGrammitis.
This species is an interesting element in the
evolution of Cochlidium, Grammitis and
StenoGrammitis. It resembles other species of
Cochlidium, particularly those species with
superficial sori, but it differs from them by
having clathrate rhizome scales, thin and brown
blade margin and markedly oblique sori. It also
resembles Grammitis by its simple blade and
inclined sori to the costa, but it differs by its
small and clathrate rhizome scales, hydathodes
present (vs. absent) and thinner (0.1 mm vs. 0.3-
0.5 mm) and brown (vs. blackish) blade margin.
More studies are necessary to delimit the genera
or to discover if more taxa are present.
ACKNOWLEDGMENTS
I thank to Carlos O. Morales (USJ) for translating
the diagnosis to Latin, the curators of the
herbaria of the Instituto de Ciencias Naturales-
Universidad Nacional de Colombia (COL),
the Museo Nacional de Costa Rica (CR), The
Missouri Botanical Garden (MO) and the
National Museum of Natural History Herbarium
(US) for their authorization to review their
specimens, to Sergio Villegas for the drawings
and the anonymous reviewers for their helpful
suggestions.
REFERENCES
Bishop LE. 1977. The American species of Grammitis sect.
Grammitis. American Fern Journal, 67: 101-106.
&&&&&&&&[  ]
Copeland EB. 1952. Grammitis. Philippine Journal of
Science, 80 (2): 93-276, t. 1-6.
&&&&&&&&[  ]
Labiak PH 2011. StenoGrammitis, a new genus of grammitid
ferns segregated from Lellingeria (Polypodiaceae).
Brittonia, 63 (1): 139-149.
&&&&&&&&[  ]
Maxon WR. 1915. Polypodium marginellum and its
inmediate allies. Bulletin of the Torrey Botanical Club,
42 (4): 219-225.
&&&&&&&&[  ]
Moran RC, Smith AR. 1995. Grammitis. In: Moran
RC, Riba R, editors. Flora Mesoamericana. Volumen
1. Psilotaceae a Salviniaceae. M&xico D. F. (M&xico):
Universidad Nacional Aut&noma de M&xico. p. 375-376.
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