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Two new species of Blechnum (Blechnaceae) from the neotropics

Alexander Fco. Rojas-Alvarado

Rojas-Alvarado, A. F. (Jardín Botánico Lankester-Universidad de Costa Rica-

Apdo. 1031-7050, Cartago, Costa Rica; e-mail: afrojasa@yahoo.com). Two new
species of Blechnum (Blechnaceae) from the neotropics. Brittonia 58: 000–000.
2006. Blechnum moranianum is a new species similar to B. loxense, and B. nigrum
is segregated from the B. fragile complex. Both are described and illustrated as result
of the author’s taxonomic fern work in Costa Rica.
Key words: Blechnaceae, Blechnum, neotropics, ferns, new species, Pteridophyta.

Rojas-Alvarado, A. F. (Jardín Botánico Lankester-Universidad de Costa Rica-

Apdo. 1031-7050, Cartago, Costa Rica; e-mail: afrojasa@yahoo.com). Two new
species of Blechnum (Blechnaceae) from the neotropics. Brittonia 58: 000–000.
2006. Blechnum moranianum es una especie nueva similar a B. loxense, y B. ni-
grum es segregada del complejo B. fragile. Ambas son aquí descritas e ilustradas
como resultado de los trabajos taxonómicos del autor en los helechos de Costa Rica.

Moran (1995) pointed out that Smith 2084
(MO) differed from Blechnum loxense
(Kunth) Hook. ex Salomon by its papillose
rachis, patent medial pinna, retroflexed basal
pinnae, dark rachis and stipe, and absence of
aerophores. This paper confirms these differ-
ences and reports others based on the exami-
nation of more specimens.
Blechnum fragile (Liebm.) C. V. Morton &
Lellinger (Lectotype, chosen by Smith, 1981:
Mexico. Veracruz: Barranca de Huitamalco,
Liebmann s.n. [Pl. Mex. 2321, Fl. Mex. 773]
(C; isolectotypes K-n.v., NY-n.v., fragm.
US!)) was originally described as Lomaria
fragile Liebm. in 1829, a genus characterized
by dimorphic fronds that was later trans-
ferred to Blechnum. Tryon and Stolze (1993)
combined this species as a variety of B. bin-
ervatum (Poir.) C. V. Morton & Lellinger,
under which they also included B. fragile and
B. ensiforme (Liebm.) C. Chr. ]. Smith
(1995) considered B. binervatum indistinct
from B. ensiforme. Several authors (Stolze,
1981; Proctor, 1985, 1989, Mickel & Beitel,
1988; Mickel & Smith, 2004; Moran, 1995)
accepted B. fragile and B. ensiforme as valid
species. Rojas (2005) segregated a new
species from B. fragile based on having
shorter brown scales without a long-attenuate
Brittonia, 58(4), 2006, pp. 000–000.
© 2006, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
apex, a yellowish stipe, fewer pinna pairs,
and a different altitudinal distribution. Here,
another new species is segregated from this
group.
Blechnum moranianum A. Rojas, sp. nov.
Type: Costa Rica. San José: Pérez
Zeledón, Páramo, Villa Mills, entre La
Georgina y Las Torres, 9°34'15"N,
83°45'00"W, 3270 m, 8 Jan 2005, Rojas &
Frias 6345 (holotype: CR; isotypes: F,
K, MO, UC, US, USJ). (Fig. 1A)
Species Blechno loxensi (Kunth) Hieron. similis,
foliis sterilibus fertilibusque dimorphis, illis sterilibus ar-
cuatis, laminis ellipticis vel elliptico-oblongis, pinnis
sterilibus mediocre involutis, fertilibus non involutis axe
atrobrunneo, pinnis basalibus retroflexis, ceteris ad
rachim perpendicularibus, aerophoris instructa recedit.
Terrestrial; rhizome 7–10 mm diam., erect,
not arborescent, not stoloniferous, the scales
10–20 × 2–5 mm, ovate to lanceolate, con-
colorous, light brown to brownish yellow, en-
tire; sterile and fertile fronds dimorphic, the
sterile ones curving to involute, the fertile
erect and skeletonized; sterile fronds 25–70
cm long; stipe 4.5–40.0 cm long, basally
stramineous to atropurpureous, the scales
3–15 × 1–3 mm, lanceolate, yellowish, with –
an entire to short-ciliate margin; blade 20–35 –
–
ISSUED: 00 Month 2006
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2 brittonia
Fig. 1. A. Holotype of Blechnum moranianum (Rojas & Frias 6345, CR). B. Representative specimen of B.
loxense (Rojas & Coto 2765, CR).
× 5.5–11.5 cm, pinnate, elliptic to oblong-
elliptic, slightly to strongly reduced at the
base, without vestigial pinnae, the terminal
pinna similar in form to the lateral pinnae;
sterile pinnae 1.5–6 × 0.5–1.0 cm, 22–26
pairs, the basal ones commonly retroflex, the
medial and distal ones perpendicular to the
rachis, sometimes falcate at apex, strongly
involute, sessile but not adnate, the margins
entire, the veins impressed; rachis without
papillae, stramineous to brownish yellow,
densely scaly, the scales 1–5 × 0.5–2.0 mm,
stellate to lanceolate, yellowish, with dentate
to ciliate margins; costa stramineus, densely
scaly, the scales 1–3 mm long, similar to the
rachis scales; fertile fronds a little longer
than the sterile; stipe (10–) 20–45 cm long;
fertile pinna 3.0–5.5 × 0.2–0.4 cm, dark
s– brown adaxially, arching at the apex;
o– aerophores present.
l–
[VOL. 58
Distribution.—Costa Rica, Cordillera Cen-
tral and Cordillera de Salamanca, from 2800
to 3270 m.
Etymology.—This species is dedicated to
Robbin C. Moran, Curator at The New York
Botanical Garden, in recognition of his many
contributions to the study of ferns and be-
cause he first pointed out the differences be-
tween this new species and Blechnum lox-
ense.
This species differs from Blechnum lox-
ense by dimorphic (vs. subdimorphic) sterile
and fertile fronds, the sterile ones arching
(vs. erect), elliptic to oblong-elliptic (vs.
ovate to oblong) blades, sterile medial pinna
involute (vs. strongly involute), fertile pinna
not involute (vs. involute), dark brown (vs.
green) adaxial lamina surface, basal pinna
retroflexed and the others perpendicular to
the rachis (vs. all ascending), and aerophores
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2006] rojas-alvarado: blechnum (blechnaceae) 3

Table I
Comparison of three species of Blechnum in Costa Rica

Characteristic B. loxense B. moranianum B. sessilifolium

Frond dimorphism subdimorphic dimorphic dimorphic

Sterile frond large 25–55 cm 25–70 cm 70–120 cm long
Sterile frond position ascending arching, horizontal arching, horizontal
Sterile blade form ovate to oblong elliptic to oblong-ellip- oblong-lanceolate
tic
Pinna pair 18–25 22–26 27–70
Basal sterile pinna ascending retroflexed perpendicular
position
Medial sterile pinna 2.5–6 cm 1.5–6 cm 6–10 cm
large
Sterile pinna margins strongly involute (abaxial involute (abaxial surface involute (abaxial surface
surface not visible) visible) visible)
Fertile pinna adaxial green brown to black brown to black
color
Aerophores absent present present
Buds absent absent present in the axes of
lateral pinna

Fig. 2. Comparison of growth habits. A. Blechnum moranianum. B. Blechnum loxense.

present (vs. absent). It differs from B. sessili-
folium (Klotzsch ex H. Christ) C. Chr. by
shorter (25–70 cm vs. 70–120 cm) sterile
fronds, elliptic to oblong-elliptic (vs. oblong-
lanceolate) sterile lamina, 22–26 (vs. 27–70)
pinna pairs, the basal pinnae retroflexed (vs.
perpendicular), shorter (1.5–6 cm vs. 6.10
cm) medial pinnae, and absent (vs. present)
buds. (Table I; Fig. 1). –
In páramo habitat at the type locality, –
–
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4 brittonia [VOL. 58

Fig. 3. Holotype of Blechnum nigrum (Smith & Vásquez 4898, UC).

s–
o–
l–
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2006] rojas-alvarado: blechnum (blechnaceae) 5

Table II
Comparison of three species of Blechnum in the neotropics

Characteristic B. fragile B. fuscosquamosum B. nigrum

Rhizome scale density dense (rhizome not dense (rhizome not sparse (rhizome visible)
visible) visible)
Length of longer rhizome 8–15 mm 5–10 mm 5–10 mm
scales
Width of rhizome scales 0.3–0.5 mm 1–2 mm 0.5–1.0 mm
Color of the center of rhi- brownish yellow brown to dark brown blackish
zome scales (longer)
Marginal color of the yellowish brown brown to atropurpureous
longer rhizome scales
Apex of the rhizome scales long attenuate attenuate long attenuate
Color of ventral stipe and atropurpureous yellowish to brown atropurpureous
rachis
Sterile blade form linear lanceolate to linear- linear-lanceolate
lanceolate
Pinna pair numbers (35–) 40–80 pinna 25–35 pinna pairs 30–35 (–45) pinna pairs
pairs (to 50 pinna pairs
on South America
Angle between pinna and 60–80 angle degree 80–90 degree 40–50 degree
rachis with the rachis
Sterile pinna large 2.8–5.0 cm long 4–6.5 cm long 4–10 cm long
Fertile pinna wide 1.0–1.5 (–2.0) mm 1.5–3.0 mm wide Ca. 2 mm wide
wide
Altitudinal distribution 800–2100 m 2200–3200 m 2000–3300 m
Geographic distribution1 Mex, CA, Col, Ec, CR, Pan, Col, Ec, Pe, Bol
Ven, Guy, Ant Pe, Bol, Ven
1
Geographic abbreviations: Mexico (Mex), Central America (CA), Costa Rica (CR), Panama (Pan), Colombia
(Col), Ecuador (Ec), Peru (Pe), Boliva (Bol), Venezuela (Ven), Guyanas (Guy), Antilles (Ant)

Blechnum moranianum was growing with B.
loxense (Rojas & Frias 6344, CR). Blechnum
moranianum was more common; no hybrids
were found between them. The two species
differ in growth habit, allowing them to be
distinguished at a distance. Blechnum mora-
nianum has arching sterile fronds and nar-
rower brown fertile pinnae, wheras B. lox-
ense has erect sterile fronds and wider green
fertile pinnae (Fig. 2).
Additional specimens examined. COSTA RICA.
Heredia: Volcán Barba, 2800 m, 29 Jul 1926, Valerio
s.n. (CR). San José: Dota, Copey, Ojo de Agua, 0.5–3.0
km del cruce rumbo a Providencia, 9°33'24"N,
83°51'35"W, 3000 m, 24 Aug 2004, Rojas & Gómez
5895 (CR, K, MO); road from Cartago to San Isidro del
General (Pan American Hwy, Rt. 2), Km 72–73, or
21–22 km SE of El Empalme, 2800 m, 30 Jan 1986,
Smith & Béliz 2084 (CR, MO, UC).
Blechnum nigrum A. Rojas, sp. nov. Type:
Peru. Amazonas: Bongará, Shillac, N by
trail from Pedro Ruiz, 5°49'S, 78°01'W,
2300 m, 31 Aug–2 Sep 1983, Smith &
Vásquez 4898 (holotype: UC). (Fig. 3)
A Blechno fragili rhizomatis squamis angustioribus
nigrescentibus sparcis, pinnis sterilibus longioribus pro-
fundus falcatis divergens.
Hemiepiphytic; rhizome 4–8 mm in wide,
long creeping; rhizome scales sparse (rhi-
zome visible), in two types, one 5–10 ×
0.3–0.5 mm, linear, tricolorous, the central
band dark brown to black, the median ribbon
brownish yellow, the margins atropurpureous
and toothed, the teeth simple to bifurcate, the
apex long attenuate, the other scales type
0.3–1 (–3) mm long, ovate to lanceolate, bi-
colorous, the central band brownish yellow
with margins brown to atropurpureous; fronds
dimorphic; sterile fronds 50–80 cm long;
stipe 12–20 cm long (1/4–1/5 of the frond
length), adaxially yellowish, abaxially atrop-
urpureous; blade 40–65 × 6–15 cm, narrowly
elliptic to linear-elliptic, pinnatisect, glabrous
on both surfaces, gradually reduced at both
ends, with 1–3 semicircular pairs of auricules
at the base, abaxially atropurpureous,
brownish-yellow adaxially; 30–35 pinna –
pairs, ascending, arching at 40–50° angle to –
–
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6 brittonia
Fig. 4. Rhizomes and scales. A, D. Blechnum
nigrum (Smith & Vásquez 4898, UC). B, E. B.
fuscosquamosum (Kessler 8993, UC). C, F. B. fragile
(Herrera 1904, CR).
the costa, medial pinnae 4–10 × 1–2 cm,
lanceolate, not involute, entire; costa yellow-
ish, glabrous; without aerophores; fertile
fronds not seen.
Distribution.—Andes of Peru and Bolivia,
from 2000 to 3300 m.
Etymology.—The specific epithet of the
new species refers to the rhizome scales with
a black center and atropurpureous margin,
giving the scales a dark appearance.
The new species differs from Blechnum
fragile in having sparse (vs. dense), shorter
(5–10 mm long vs. 8–15 mm), blackish (vs.
brownish yellow to brown) scales. These
scales are tricolorous with an atropurpureous
margin, a yellowish line and dark brown to
black center (vs. bicolorous with yellowish
margin and brownish-yellow center). The rhi-
zome scales are mixed with smaller ones
only 0.3–1(–2) mm long, which the other two
species of Blechnum do not have. The sterile
s– pinnae consist of 30–35 pairs (vs. (35–)
o– 40–80) and are relatively longer (4–10 cm
l–
[VOL. 58
long vs. 2.8–5 cm). Blechnum nigrum is gen-
erally found at higher elevations compared to
B. fragile (2000–3300 vs. 800–2100 m).
From Blechnum fuscosquamosum A.
Rojas, B. nigrum differs by sparse (vs. dense)
and tricolorous (vs. concolorous or bicolor-
ous) scales, abaxially atropurpureous (vs.
yellowish) stipes, and more arching (40–50°
vs. 80–90°) sterile pinnae (Table II; Figs. 3
and 4).
Additional specimens examined. PERU. Caja-
marca: San Ignacio, San José de Lourdes, Campamento
Zural, camino al Cerro Picorana, 4°59'25"S, 78°54'5"W,
2010 m, 28 Jan 1999, Díaz et al. 10544 (UC).
BOLIVIA. Cochabamba: Chapare, Paractí, ca. 83
km from Cochabamba on road to Villa Tunari, ca. 6700
ft. [2045 m], 16 Nov 1979, Foster 79-190 (UC). Yun-
gas: 1890, Bang 435 (UC); Unduavi, 3300 m, Nov
1910, Rosenstock 46 (UC).
Acknowledgments
I thank Franco Pupulin for the translation
of the diagnosis to Latin, and curators of the
herbaria at the Museo Nacional de Costa
Rica (CR), Missouri Botanical Garden (MO),
and University of California (UC) for speci-
men loans, and to the anonymous revisers for
their helpful suggestions.
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——— & A. R. Smith. 2004. Pteridophytes of Mexico.
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Moran, R. C. 1995. Blechnaceae. Pp. 325–333. In:
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