Typification and Relationships of Cheilanthes incisa (Pteridaceae)

2007·4.9 MB·English

by  Mónica Ponce

American Kern Journal 97(3):14()-148 (2007)
and Cheilanthes
Relationships of
Typification
incisa (Pteridaceae)
Monica Ponce
San Argentina
Instituto de Botanica Darwinion, C.C. 22, 1642, Isidro,
Prado
Jefferson
[nstituto do Botanica, C.P. 3005, CEP 01061-970, Sao Paulo, SP, Brazil
Makta
Morhelli
Bosque
Facultad de Ciencias Naturales Museo, Univorsidad Nacional do La Plata, Paseo del
y
mOO,
La Argentina
s/nro., Plata,
—
was
Abstract. Cheilantlws incisa, small and rare endemic fern of Rio do Janeiro State (Brazil),
a
taxonomic
described in 1859. Hypolepis serrata, published ten years later, is usually considered a
synonym. Botb names have typification problems, addressed here by designation of a neotype for
among
C. incisa and selection of a lectotype of II. serrata from Glaziou materials cited by Fee.
Epidermal characteristics and spores of C. incise are described for the first time and their
diagnostic value assessed. Taxonomic relationships between C. incisa and putatively related
ehoilanlhoid ferns are also discussed.
Cone
Southern
During preparation of the pteridophyte Catalog of the of
we
South America detected taxonomic and typification problems involving
Kunze ex and Hypolepis According
Cheilanthes incisa Mott. serrata Fee. to
endemic
Mynssen & Windisch Cheilanthes incisa small fern to Rio
(2002), a
is
de Janeiro State, Brazil. Hypolepis serrata, described by Fee (1869) in the
and was
Cryptogames vasculaire du Bresil very similar morphologically
v. 1, is
names
synonym by Baker Both have
considered of incisa (1872).
a C.
The mentions one specimen
typification problems. protologue of C. incisa
now The
(without collector and from Serra da Estrella, Rio de Janeiro.
lost)
original description of H. serrata cited two syntypes [Glaziou 929 and 2336)
name.
and requires selection of a lectotype to stabilize application of the
According Mettenius Cheilanthes incisa closely related to C.
to (1859),
is
Kunze
(Hook.) Mett. and schimperi because three species
californica C. all
have dentate segments. Most recent treatments (Lellinger, 1968; Smith, 1975;
Mickel and Smith, 2004) have assigned californica Aspidotis (Nutt ex
C. to
USA
Hook.) Copel., small genus confined of the western and Mexico.
a
Aspidotis can be recognized by dentate-mucronate segments and striate
and
lamina. Cheilanthes schimperi has a paleotropical distribution (Africa
Madagascar) and was included Aspidotis by Pichi-Sermolli (1950)
in
it
because of similar morphology.
its
we and Hypolepis
paper review the typification of Cheilanthes incisa
In this
serrata and discuss the similarities and differences between C. incisa and
related species based on the morphology of the sporophyte and spores.
PONCE AND
ET TYPIFICATION RELATIONSHIPS OF CHEILANTHES
AL.: INCISA
141
Materials and Meti
iods
Hypolep
examined HBG,
materials from B, K, LE, R, RB, and The following
P.
MO
from
K.
m
Aspidotis
californica (Hook.) Nutt. ex Copel.
UNITED
STATES. mo
Specimens studied.—
Califor
mbankment
Macabes
of Creek, 1300 19 Feb 1980. M.
ca.
ft.,
Tehama
(MO);
mi
5
Red
north-northeast of occasional
Bluff, in crevices in basalt outcrops
in
Quercus May
wislizenii-Pinus sabiniana woodland, Oswald
9 1991, H.
V.
8r L.
N
Ahart 4564 (MO); Butte County, along Feather mi
trail to Falls, ca. 5 of the
May
m, 28
M. Baer 4023 (MO)
M.
(MO);
Santa Catalina Island, Avalon, canon, 1897, B. Trask 5 (MO)
densa
Aspidotis
(Brack.) Lellinger
UNITED
Specimens studied.— STATES. Apr Brown
California: 10 1934,
L. L.
1
(MO); Tulare County, Sequoia National Park, between Franklin Pass and Kern
among
8400
River, crevices granite boulders, 23
ft., trail, Jul 1942, B. S. Ferris
8r L. Lorraine 10792 (MO); Del Norte County, Mill Creek, 27 Jun-1 1922,
Jul L.
Abrams
B. 8423 (MO); Mis. about the head waters of Sacramento River, rocky
3Sep
2778
places, 1882, Pringle (MO).
C. G.
Aspidotis
meifolia Eaton) Serm.
(D. C. Pic.
—
MEXICO. San
Specimens studied. Luis Potosi: near Los Canos, 15-21 Oct 1902,
Palmer 264 (MO); Nuevo Madre
E. Leon: Sierra above Monterrey, 3000
ft.,
shaded Aug
banks, 8 Jun 1888, C. G. Pringle 1987 (MO), 27 1903, C. G. Pringle
11778 (MO, Coahuila: Mun. Muzquiz, Rancho Agua wooded
SI); Dulce,
canyon on
the east slope of the Sierra de San Manuel, 28 Jun 1936, Lyle
F.
Wynd
H. Mueller 350 (MO). Tamaulipas, oak on Jaumave
8- C. road about
forest
13 miles southwest of Ciudad rough limestone mountainsides
Victoria, near
summits May
of Sierra Madre, ca. 100 m, abundant, 13 1949, B. McVaugh
10516 (MO).
Cheilanthes Kunze
incisa ex
Mett.
—
Man
BRAZIL.
Specimens Rio de
studied. Janeiro:
ombrio limido na mata
e
J.M.
Rio de 12289
laneiro, Jan 1881, A. Glaziou
17929
Ule (UC); "Brasilia", Biedel 1822
(K).
VOLUME NUMBER
AMERICAN
FERN JOURNAL:
97 (2007)
142 3
Kunze
Cheilanthes schimperi
MALAWI.
Specimens studied.— Southern, Mi. Mulanjo foot Likabula Valley,
-
cm mass
up rhizome surrounded by dense of
900 30 a
in, little fern, to tall,
dimorpha,
on shady road bank, nearby are Actiniopteris
localized
roots,
Chapman
&E. 9498
Adiantum phillipense, Pellaea spp., 20 Feb 1989, D. G.
/.
Rum
escarpment, 1000 m,
(MO); North Region, phi Livingstonia
District,
DEMOCRATIC
3594 (MO).
ix
Mabondo
Meridional) Riviere
VI
Apr Bodei Malai a (Shaba
4 1987, A.
galerie,
m
Boden-
Meridional) Kintu 27 28'E 11 49\S 1250 Termiliere lFeb 1985 A.
& ZAMBIA. Mazabuka
ghien Malaisse 46 (MO). Southern Prov., Distr.,
F.
Kafue Gorge below Kafue Dam, 15 48'S, 28 25'E, 950 m, humid, shaded slope
Komds
woodland, 30 Dec 1971, 6741 (MO).
in
/.
—
NaOH
and
Morphological Several Fronds were bleached with (5%)
studies.
mounted The
and
sodium hypochlorite solution (5%), stained in glycerin jelly.
observations and illustrations were made using the light microscope Wild
M5
M2() and stereomicroscope Willd both with camera lucida drawing.
—
Spores. Spores obtained from dry herbarium specimens in the list were
studied by microscopy (LM) and scanning electron microscopy (SEM).
light
LM 3%
sodium min and
For the spores were treated with hot carbonate for 2
SEM
acetolysed according to Erdtman (1960). For the material was treated with
3%
sodium carbonate min, washed with distilled water, dehydrated,
hot for 2
96%
suspended ethanol, and then transferred to acetate plates. After drying
in
were performed
these were coated with gold. All of the observations witli
microscope and JEOL JSMT-100 scanning electron microscope
a BFI2 light a at
Mnseo
he Faoultad de Cieneias Naturales v de La Plata.
I
Results
—
HBG,
Typificalion. The syntypes of Hypolepis serrata were located at B, K,
and At K, among the specimens identified and cited by Baker in the Flora
P.
we two one
found one specimen [Glaziou 2336) with labels:
Brasiliensis,
name and another with For Baker, the
containing the serrata C. incisa. latter
II.
name
was There an isosyntype, Glaziou
epithet the correct for this material. is
HBG we we BM.
photograph
could study only located in
929, in but not a
it;
we
Based on information, chose the authentic specimen, Glaziou 2336,
this
we K and Hypolepis
had seen and studied as the lectotype of serrata.
that at P,
No any annotated
authentic type material of Cheilanthes incisa or other old
Kunze were found European
specimen by Mettenius the herbaria.
or in
According the TL-2 (Stafleu and Cowan, 1981) the types of species described
to
LZ Second
by Mettenius and Kunze housed were destroyed during the
at
we which
from
World War. Thus, for Cheilanthes incisa selected a neotype, is
summary
The taxonomic
same and Glaziou.
the area district as the original is
as follows.
AND
PONCE ET TYPIFICATION RELATIONSHIPS OF CHEILANTHES INCISA 143
AL.:
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BOTANICO DO DE JANEIRO
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Fam. .^icm.<i.^ceju
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Uktl^k^-lki^ I.MCLA& £x....HtTt.:
Ao
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Proced.. Z>a(- ^.Q.........J..At/H.4ajAO.£"
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Col.
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Data
/3
, .
At Lectotype of Hypolepis serrata Fee, Glaziou 2336 (K). At bottom. Neotype of
top.
Fig. 1.
Cheilonthes inciso Kunze ex Mett., C. Brarfe 76255 (duplicate at SI).
/I.
VOLUME NUMHMK
^VlLRICAN FERN JOURNAL:
144 07
3 (2007)
mm
1
>\S,
E
Lamina and epidermal
Fig. 2, characteristics. A-I5. Aspidotis californica look.) Null, ex Copel.
(I
C. Aspidotis densa (Brack.) Lellinger. D-E. Aspidotis maifolia C. Eaton) Pic. Serm. F-G.
(I).
Kimze
Chailanthes incisa ex Mett. H-I. Cheilanthas schimperi
Kun/.e.
Mett
Mun
Janeiro:
800 m, nas pedras, 13 Jun 1940, A. C. Brade 16288 (RB, duplicate
at
SI).
Hypolepis
serrata
Fee,
Cr. vase. Br. 1. 53, tab. 13, fie. 3. 1869. Lectotype
J
(Fig.
1).
—
m
Aspidotis
(sensu
Smith, 1975) distinguished from most other cheilanthoid by havin
is ferns
lamina
striate surfaces. This striation formed by epidermal
cells that are
is
and
elongate with thickened These
parallel walls. aspects can be observed
in
O
z
>
id
—
Table Comparative spore characteristics of the studied taxa. n
1.
>
Cheilanthes Aspidotis Aspidotis Aspidotis Cheilanthes G
mei schimperi
mcisa densa folia
californica
>
Characteristics a
globose globose
Shape subglobose, tetrahedral- subglobose globose
globose
>
brown brown, brown brown brown
Color yellow dark light light light
light
um
Equatorial diam. 34-35 |im 51-60 |im 43-45 |xm 32-46 40-48 |im 5
(
Polar diam. 27-34 |im 45-48 |im 40-50 |xm 34-39 |im 36-58 |jm
en
1-3 2-3 3-4.5
Perispore thickness 1-2 \xm 2-3.5 jam urn jam Jim
CO
Perispore Structure
O
Surface rugulate, rugulate-cristate rugulate-cristate rugulate-cristate reticulate, reticulate- cristate-reticulate
cristate
Cristae 0.5 jam, isolated 1.5 |im 2 |im 1-1.5 |im |im
continuous
Outer stratum continuous continuous continuous discontinues
Middle stratum interwoven threads interwoven threads interwoven threads interwoven threads interwoven trabeculae
Inner stratum thin, continuous thin, continuous thin, continuous thin, continuous thin, continuous
CO
Q
CO
AMERICAN FERN VOLUME NUMBER
140 JOURNAL:
97
(2007)
3
Fig. 3. Spores. A-D. Cheilanthes incisa Kunze ex Mett. A. Distal view. Equatorial view.
B. C.
Fracture of the sporodermis. D. Detail of the surface, Braga et al.4427 (RB). E-G. Aspidotis
californica (Hook.) Null, ex Copel. E. Distal view. F. Fracture of the sporodermis. C. Detail of the
&
Oswald
surface, Ahart 4564 (MO). Aspidotis densa and
H-J. (Brack.) Lellinger. H. Distal
proximal view. Equatorial view.
Detail of the surface, Ferris Lorraine 10792 (MO). K-M.
I. Er
J
M
Aspidotis
meifolia (D. C. Eaton) Pic. Serai. K. Distal view. Fracture of the sporodermis.
L. Detail
Wynd
of the surface, fr Mueller 350 (MO). N-O. Cheilanthes schimperi Kunze. N. Distal view. O.
&
Fracture of the sporodermis. P. Detail of the surface, Bodenghieu Muhalci 2324 (MO).
AND
PONCE ET TYPIFICATION RELATIONSHIPS OF CHEILANTHES INCISA 147
AL.:
A. californica (Hook.) Nutt. ex CopeL, A. densa (Brack.) Lellinger, and A.
meifolia (D. C. Eaton) Pic. Serm.(Fig.
2).
Although superficially similar to Aspidotis in the shape of the dentate-
and
mucronate ultimate segments, Cheilanthes incisa schimperi differ in
C.
having smooth laminae with sinuous, thin-walled epidermal
(non-striate) cells
and by the with lobate-laciniate pseudoindusia having erose or dentate
sori
margins Tryon and Tryon (1982) stated that the apparent morpholog-
(Fig.
2).
Group" was
ical similarity of C. schimperi to the rest of their "C. californica
due morphological convergence, not true relatedness, and they excluded
to to
schimperi from the group.
C.
Other characteristics that distinguish Cheilanthes incisa from Aspidotis and
schimperi rhizome and chartaceous subcoriaceous lamina
are erect or
C. its
with hidden venation. Cheilanthes incisa occurs isolated in southeastern
and no from and South America morpholog-
Brazil other species Central
is
ically similar to
it.
and by
Cheilanthes schimperi from incisa Aspidotis species
differs C. its
nodose rhizomes, membranaceous lamina with very thin mesophyll, and
easily visible venation.
The and
spores of Aspidotis Cheilanthes studied here are similar to the other
cheilanthoid ferns such as Argyrochosma and Pellaea. The perispore with
three layers the main feature of the group (Morbelli and Michelena, 1989;
is
The
Morbelli and Ponce, 1997; Morbelli 2001). spore characteristics are
et
a/.,
SEMs shown The
summarized and
in Table of the spores are in Fig.
1 3.
studied material had some well-formed spores, but collapsed spores and
between and monolete were
spores intermediate frequent.
trilete
Discussion
between dentate-mucronate lamina Cheilanthes
Despite the of
similarities
incisa, C. schimperi and Aspidotis, the former species lack the striate leaf
and
tissue (formed by elongate epidermal cells with their thickened walls)
Our
well-defined, scarious indusia characteristic of the genus. studies
latter
showed from
that the spores of incisa are slightly different those of
C.
Aspidotis and schimperi as well. Although the palynological characteristics
C.
of the studied species are those of Cheilanthes, this genus highly
is
we
polymorphic Smith and uncertain
(and polyphyletic; ah, 2006), are
et
whether the observed spore features are indicative of a close relationship.
we
Based on the morphological evidence here presented retain Cheilanthes
incisa and schimperi in Cheilanthes. However, the final decision regarding
C.
placement
the correct generic of this species requires further study.
Literature Cited
and
Baker, G. 1870. Cyatheaceae et Polypodiaceae. Pp. 305-623, in Martius, C. F. P. A. G. Eichler,
}.
eds. Flora Brasiliensis v.l, part Fleischor, Leipzig.
2.
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148 97
3 (2007)
A
The
Ekdtman, 1000. acetolysis method. revised description. Svensk Botanisk Tidskrift
Ci.
54:501-504.
F£e, A. L. A. 1809. Cryoptogames vasculaires du Bresil v. 1. Veuve Berger- Levrauuld & Files, Paris.
A
Lellinckr, D. 1908. note on Aspidotis. Amer. Fern 58:140-141.
B.
].
»
Uher
Mettenius, 1859. einige Farngattungen. Bronner, Frankfurt.
(I. II. L.
M. and
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I.
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J.
Morbelli, M. A., M. M. Ponce, C. MacIck and M. R. Pineiro. 2001. Palynology of Soulh American
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Smith, A. R. 1975. California species of Aspidotis. 23:15-24.
M. A
Smith, A. R., K. Pryer, E. Schukttpelz, P. Korall, H. Schneider and P. G. Wolf. 2000.
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nd
Stakleu, F. A. and R. S. Cowan. 1981. Taxonomic Literature 2 edition. Vol. Lh-O. Bohn.
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