Melpomene anazalea, a New Species of Grammitid Fern (Polypodiaceae)

2008·3.2 MB·English

by  Michael Sundue

New
Melpomene
Grammitid
anazalea,
a Species of
Fern
(Polypodiaceae)
Marcus 1
Lehnert
itut fur Pflanzenwissenschaften, Abt. Systematische Botanik, Geoi
Gottingen, Untere Karspule D-37073 Gottingen, Germany
t 2,
elpomene anazalea Sundue & Lehnert from
ted rock crevices in relatively dry paramos.
The
grammitid
ferns are a monophyletic group within Polypodiaceae
the
-
(Schneider et al., 2004) and comprise about 750 species with a pantropical
southern temperate They
distribution (Parris, 2003). are predominantly small
plants exhibiting epiphytic and saxicolous and
habits Kubitzki
(Parris in
Green, Moran
1990; Parris 2001, 2003; Smith and Melpomene
in Riba, 1995).
is
characterized by having brown
ventral root insertion, reddish
clathrate scales
with and
sometimes
apical also marginal gland-like hydathodes without
cells,
calcareous and
deposits, distinguished from other grammitid genera by
is
having secondary
metabolites with a distinctive sweet and spicy odor (Smith
and
Moran, While
1992). the circumscription of several genera within the
grammitid has been
ferns called into question by the phylogenetic analysis of
Ranker
et al. (2004), further studies continue to support the monophyly of
Melpomene
Many
(Lehnert
et in press). recent and herbarium
al., field trips
studies have yielded more new
species
science
to (Labiak, 2000; Rojas, 2001)
one MS
like the presented which was
here, discovered by while conducting
work
field in Colombia.
Terminology
follows
that of Lellinger Grammitid with
(2002). ferns are beset
two
types of laminar indument
that appear be and
to distinct not part of a
homologous
The
term
series. "seta" has been
often applied the
to
stiff,
darkened
hairs that frequently occur in grammitid Melpomene
ferns including
(Smith and Moran,
1992; Labiak and Prado, The same
2005). treatments used
the term
"hairs" for the paler, often branched trichomes found mainly on
young
and
petioles rachises Melpomene. we
in However,
find that variation
occurs within
the "setae" Melpomene,
of sometimes
representing species
address:
Staatliches
t
SUNDUE & LEHNERT: MELPOMENE ANAZALEA, A NEW SPECIES OF GRAMMITID FERN
209
autapomorphies and sometimes growth
reflecting different conditions. For
example, the "setae" of M. firma Sm.) A. R. Sm. & R. C. Moran are usually
(J.
quite short and patent. For this condition the term "acicular hairs" would be
more fitting. In M. flabelliformis (Poir.) A. R. Sm. & R. C. Moran and many
related species, the "setae" are sometimes flaccid and twisted. Such trichomes
are better described as "ciliform hairs" (Lellinger, 2002). For the straight
stiff,
trichomes term
or "setae", the "setiform hairs" available (Lellinger, 2002).
is
Furthermore, used in Latin diagnoses, the term "seta" would translate as
which
"bristle" (Steam, 2004), a notably different structure (Lellinger, 2002)
is
We
Melpomene
that not found in or other grammitid genera. prefer the terms
is
and
"acicular," "ciliform," "setiform hairs" describe the simple pericel-
to
lular trichomes that are usually dark red in color with acute and "clavate
tips,
hairs" for the minute simple and branched, pericellular hairs that are pale in
and
color bear clavate cells at their apices or along their length. This second
type of hair has been referred to as "glandular" (Parris, 2005) and appears to be
homologous
to the secretive gland-bearing branched hairs of Zygophlebia
we
(Bishop, 1989), but in the course of our studies, have not observed a
Melpomene.
secretive nature in the hairs of
—
Melpomene TYPE COLOMBIA.
anazalea Sundue &
Lehnert, nov.
sp.
SFF
Boyaca: Municipio Chisaca-San Pedro de Iguaque, de Iguaque, sendero
Apr
subiendo a laguna de Iguaque, 05°41'3.1"N, 73°26'22.2"W, 3536 m, 30
la
& HUA,
Sundue NY;
2007, A. Vasco 1290 (holotype: isotypes: UC). Figs.
1, 2.
A
Melpomene peruviana (Desv.) A. R. Sm. & R. C. Moran statura minore
cm cm
X X M.
(laminis 1.0-3.7 0.2-0.3 1.5-8.5(9.0) 0.4-1 in peruviana),
vs.
pilis setiformibus in pagina superiore crebris (vs. absentibus), in pagina
inferiore aequaliter distributis (vs. in soris confertis) differt.
Plants saxicolous, growing in moss layers and rock crevices (Fig.
1).
Rhizomes short-creeping, horizontal 2A), frequently branched, 0.4-
(Fig.
mm
cm
0.5 diam. Fronds 1.3-4.7 long, appressed to substrate, inserted onto
the rhizome at acute to right angles, closely to moderately spaced (1.5-
mm,
mm). 3-4 X 6-10 wide, dark brown,
3.0 Scales 0.3-0.4 cells clathrate,
apex
iridescent, narrowly cordate the base, attenuate at the (Fig. 2B); apical
at
3-10 X
gland-like cells 1-4, linearly to furcately arranged (Fig. 2C). Petioles
0.2-0.3 mm, proximally distally marginate, with few to many, dark
terete,
mm
brown, spreading, simple and branched
setiform 1.3-1.5 long,
hairs,
mm
clavate hairs present on crosiers and persistent on older fronds, 0.2-0.4
and sunken
long 2D). Rachises castaneous to blackish, planar slightly
(Fig.
with
adaxially, hemispherically protruding abaxially, scattered setiform hairs
mm
on both the abaxial and adaxial sides, the setiform hairs 1.0-1.5 long,
provided with
spreading, dark brown, the abaxial side of the rachises also
Laminae X
scattered simple and branched clavate hairs. 1.0-3.7 0.2-0.3 cm,
linear to narrowly elliptic, the bases cuneate, the apices acute, 1-pinnatisect
throughout, with 10-20 pairs of segments; segments oblong, fully adnate, the
bases slightly decurrent, the apices rounded, the costae not visible (Figs. 2E,
1-2 proximal segment
F); pairs usually markedly smaller than subsequent
segments, and with more
the base strongly decurrent, the abaxial and adaxial
surfaces of the lamina as well as the segment margins densely provided with
mm
brown
dark
setiform
hairs, 1.0-1.5 long, spreading 2E, the
(Figs. F),
abaxial surface of the lamina provided with and
also simple
scattered
branched
clavate hairs; stomata not hydathodes and
visible; present, small
inconspicuous; 1-3
sori per segment, confluent at maturity, with setiform hairs
like those of the laminae
(Fig. 2E).
The name
=
refers to the drought resistance of the species (Greek an-
not;
=
azaleos
dry).
This minute and known
distinctive species only from
is a single collection
made
in Santuario Flora Fauna Iguaque
y in the Cordillera Oriental of
Colombia, where grows on
it protected rock ledges in dry paramo with
No HUA
Espeletia at 3536 m. further collections were located COL.
at either or
The SFF
Iguague on
situated the western
is slope of Colombia's
Cordillera
Oriental in a region of relatively dry paramos
(Rangel-Churio, 2000). Despite
known
we
being from
a single gathering,
are confident
that this plant
new we know
represents a species from
as
a revision of the genus
(Lehnert,
unpublished
data) that no comparable specimens
are located in the major
world
herbaria of the Acknowledgements) and
(see no
that species
exist for
SUNDUE NEW
& LEH \X AZALEA, A SPECIES OF GRAMMITID FERN
:
AMERICAN FERN VOLUME NUMBER
JOURNAL:
212 98 4 (2008)
which M. anazalea could be mistaken for or regarded as a mere morphotype.
It
km
which
remarkable that this species, occurs within 130 of Bogota, has not
is
been collected and brought to our attention until now.
Melpomene many
anazalea most M. peruviana
in respects similar to
is
&
(Desv.) A. R. Sm. R. C. Moran, but differs from that species by having
abundant setiform hairs on both sides of the rachises and laminae that are not
By
clustered in the comparison, the laminae of M. peruviana have
sori.
The
setiform hairs clustered in the sori but are adaxially glabrous. rachises of
M. peruviana moderately and moderately on
are setose abaxially, setose the
proximal half adaxially. The segments of M. anazalea are commonly round
at
their tips, whereas those of M. peruviana are often more deltate (to 3 times
longer than broad) with more acute Melpomene peruviana
tips. further differs
by having readily visible stomata on the abaxial side of the lamina. Otherwise,
the similarities concerning morphology and ecology are considerable. The
rhizomes and scales of the two species are indistinguishable, and both occur in
areas that are apparently too dry for other members of the genus. The
may
appressed fronds be an adaptation to avoid desiccation in the cold and
windy
environment of the paramos. With nearly isodiametric segments and
its
common
its clearly alate petioles, M. anazalea also reminiscent of the
is
M.
species moniliformis (Lagasca ex Sw.) A. Sm. & Moran which was
R. R. C.
&
collected nearby {Sundue A. Vasco 1288, NY), but M. moniliformis appears
virtually glabrous at first sight (shorter, often ephemeral and inconspicuous
common
hairs are on and and
petioles in the sori) has straight, erect fronds.
uncommon
Adaxially setose laminae are in Melpomene. Other species with
Melpomene
this character are pilosissima (M. Martens & Galeotii) A. R. Sm. & R.
Moran, M. huancabambensis
C. Lehnert nov. and M. michaelis
sp. ined.,
Lehnert sp. nov. ined. These species are much larger than M. anazalea, have
all
longer segments
(3-5 times longer than and have
broad), fronds that are not
appressed
to the substrate but rather hang from branches rocky
freely tree or
Of
ledges. these species, only M. pilosissima occurs within the range of M.
anazalea, but grows lower montane
it at elevations in forests; the other two, M.
huancabambensis and M. known
michaelis, are only from the central Andes.
AAU,
of the following BM, BHCB,
s herbaria:
B, C, C(
COLO, CUZ, GOET,
K, LOJA, LPB, P, QCA, QCNE, RB, S, SP, TUR, UC, US, USM. He thanks the DA
(German Academic Exchange DFG
Service) and the (German Research Foundation) financing
for
fi
and
trips to Bolivia Ecuador in the years 2002-2004
as part of his studies on Melpomene.
MS
thanks Alejandra Vasco
Gutierez, Alejandro Marin, and
Octavio Betancur
lor assistance
Literature Cited
new
ov, L. E. 1989. Zygophlebia, a genus of Grammitida*
New
ak, P. H. 2000. species and new
combinatio:
(Pteridophyta). Brittonia 52:246-255.
SUNDUE MELPOMENE AN NEW
& LEHNERT: AZALEA, A OF GRAMMITID
SPECIES FERN
213
Labiak, P. H. and Prado. 2005. As especies de Melpomene e Micropolypodium (Grammitidaceae-
J.
Pteridophyta) no Brasil. Bol. Bot. Univ. Sao Paulo 23:51-69.
M.
Lehnert, M., Kksm.hr. A. Schmidt-Leb^hn, and
Ni. S. Klimas, S. Fehlberc; T. A. Ranker, (in press).
Phylogeny genus Melpomene DNA
of the (Polypodiaceae) from
inferred chloroplast
analysis.
Systematic Botany.
A
modern
Lellinger, D. B. 2002. multilingual glossary taxonomic
for pteridology. Pteridologia
3.
American
Fern
Society.
Parris, B. S. 1990. Grammitidaceae. In: Kubitzki, K. (ed.). The families and genera of vascular
New
plants: vol. Pteridophytes and gymnosperms. Springer Verlag, York.
1.
Taxonomy
Parris, B. S. 2001. of Malesian Grammitidaceae in relation to phytogeography and
ecology. Pp. 155-160, In: Saw, L. G., et al. (eds.). Taxonomy, the Cornerstone of Biodnrisitx
Proceedings of the Fourth International Flora Malesiana Symposium 1998.
(Forest research
Parris, B. S. 2005. Grammitidaceae. Flora of Tropical East Africa. Royal Botanic Gardens, Kew,
U.K.
Rangel-Churio, O. 2000. Clima. Pp. 85-125, Rangel-Churio, O. Colombia Diversidad
In: (ed.).
J. J.
Biotica La region de vida paramuna. Instituto de Ciencias Naturales-Instituto Alexander
III.
von Humboldt,
Bogota.
Ranker, T. A., A. R. Smith, B. S. Parris, M. O. Geicer, C. H. Haufler. S. C. K. Straub and H.
J.
Schneider. 2004. Phylogeny and evolution of grammitid ferns (Grammitidaceae): case of
a
rampant morphological homoplasy. Taxon 53:415-428.
Rojas, A. F. 2001. Seis especies y dos nuevos registros de helechos (Pteridophyta) para Costa Rica.
Rev. Trop. 49:435-452.
Biol.
Schneider, H., A. R. Smith, R. Cranfill, T. Hildebrand, C. H. Haufler and T. A. Ranker. 2004.
].
Unravelling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae):
exploring aspects of diversification of epiphytic plants. Mol. Phylo. Evol. 31:1041-1063.
Smith, A. R. 1995. 30. Grammitidaceae. Pp. 366-393, In: Moran, R. C. and R. Riba, (eds.). Flora
Mesoamericaiui. Volumen Ptondohtas, Psdotaceae a Salviniaceae. Univ. Nacional
1.
Autonoma
de Mexico, Ciudad
Universitaria.
new
Smith, A. R. and R. C. Moran. 1992. Melpomene a genus of Grammitidaceae (Pteridophyta).
th
Stearn, W. T. 2004. Botanical Latin ed.). Timber Press, Portland, Oregon, USA.
(4