Table Of ContentVolume 78
Annals
Number
of the
1991
Missouri
Botanical
Garden
MONOGRAPH
THE
OF
Robbin Moran^
C,
NEOTROPICAL FERN GENUS
STIGMATOPTERIS
(DRYOPTERIDACEAE)'
Abstract
Stigmatopteris contains 24 species with no varieties or subspecies. distinguished from other genera of
It is all
numerous
Dryopteridaceae (sensu stricto) by punctate glands in the lamina, scales with uniseriate teeth or cilia,
glandular terminating the apices of the teeth or and clavate vein apices that end behind the leaf margin.
cells cilia,
Other characteristics shared by or nearly species of the genus but also found in other dryopteroid ferns are
all all
nonindusiate sori, thin-textured leaves, glabrous laminar tissue, acuminate-serrate pinna, apices, thin, flaccid, tan or
orangish scales, erect or short-creeping stems, and minutely pubescent adaxial grooves of the axes. Within the
Dryopteridaceae, Stigmatopteris forms a clade with Cyclodium, Olfersia, and Polybotrya, held together by the
synapomorphy of pubescent adaxial grooves of the axes. The genus entirely neotropical and primarily montane,
is
Amazon
ranging from the Antilles and southern Mexico to southern Brazil but conspicuously absent from most of the
The
basin. The Andes are the center of diversity for the genus, harboring eleven species, two of which are endemic.
coastal mountains of southeastern Brazil are another center, with six species, all of which are endemic. The Greater
Antilles have also been important in the diversification of Stigmatopteris because they contain three species, all of
which are endemic. All members of the genus grow in wet forests, generally from 200 to 2,000 m.
Resumen
Stigmatopteris contiene 24 especies sin subespecies ni variedades. Es distinguido de todos los otros generos de las
dryopteridaceas (sensu stricto) por las numerosas glandulas punctatas, escamas con dientes o cilias numerosas, celulas
glandulares terminando los apices de los dientes o cilias, y apices clavatas de las venas que terminan atras del margen
de la lamina. Otras caracteristicas compartidas por todas o casi todas las especies en el genero, pero encontrado
thank Alan R. Smith (UC) for helpful discussions on all aspects of this work and Robert G. Stolze (F) for first
^ I
pointing out to me that Stigmatopteris needed a modern monograph. RoUa M. Tryon (then at GH, now at USF)
reviewed the nomenclatural parts of the species treatments and suggested improvements. Jorge V. Crisci (LP) provided
HENNIG86
(MO)
assistance in running the program used in the cladistic analysis of the species, and Enrique Forero
and Warren D. Stevens (MO) facilitated funding for annotation labels and scanning electron microscopy. Gerrit
Davidse (MO) and Henk van der WerfF (MO) also provided support and encouragement for this project. thank the
I
AAU, BM, CM, COL, CR,
curators of the following herbaria for making their specimens available for study: B,
DUKE, F, G, GH, K, L, LIL, MICH, MU, NY, P, PH, PORT, QCNE, SI, UC, US, VEN, Z.
Missouri Botanical Garden, P.O. Box 299, Louis, Missouri 63166, U.S.A.
^ St.
857-914.
Ann. Missouri Bot. Card. 78: 1991.
858 Annals of the
Garden
Missouri Botanical
tambien en otros helechos dryopteroides, son los soros no indusiados, hojas meinbranaceas, tejido de la lamina glabro,
apices de las pinnas acuminadas-seratas, escamas membranaceas. flacidas, pardas claras o naranjizas, rhizomas rectos
En
o corte-reptantes, y los surcos adaxlales de la lamina cortamente pubescentes adentro. las dryopteridaceas,
Stigmatopteris hace un grupo monofiletico con Cyclodiurriy Olfersia, y Polybotrya, basado en la sinapomorfia de
surcos adaxiales pubescentes. El genero es completamente neotropical principalmente en regiones montanosas con
y
bosques, extendiendose desde las Antillas y sur de Mexico hasta el sur de Brasil pero ausente de la mayoria de la
cuenca Amazonica. Los Andes son centro de diversidad para genero, con once especies, dos de las cuales son
el el
endemicas. Las montanas costales del sureste de Brasil son otro centro, con seis especies, todas las cuales son
endemicas. Las Antillas ban side importante tambien en la diversificacion de Stigmatopteris, con tres especies, todas
200
de las cuales son endemicas. Todas las especies en el genero existen en bosques humedos, generalmente de a
2,U0U m.
Taxonomic History diversity and endemism for the genus the Andes,
is
and two endemics
containing eleven species (S.
Stigmatopteris was proposed by Christen-
first
Two
ichthiosmdy and pellucidopunctata). spe-
5.
sen 1909. Until then, of species had been
in all its
and pterorachis) are near-
cies {S. hiilbifcra S.
placed Dryopteris, a genus that comprised a
in
Andean
^^^^^-^^^ ^, ^^ey also occur outside the
unrelated assemblage of nearly 1,000 species.
vast,
Region the Santa Marta Mountains of northern
in
Four years later, in the volume of his Dryop- ^ u n ^u\.
first
^ ' Col, omb,ia. . Wvyi/-t.uh-m *tuhe Aandiean Kegi•on, Loi\ ombia
^^' u-
monograph, Christensen (1913)^ c1hanged1 his a
terts ^
^ andj iL?cuadjor uhave more speci-es /(*ten)\ Xm any
. . ,
mind about the rank of Stigmatopteris, placing
it
other country (Tables
1, 2).
back Dryopteris as a subgenus, but with the
in
The mountains northern Venezuela
coastal of
-re-
most
proviso that could be treated, alone with
u
it r u-
*^ J- M ,have ^two species ot otigmat*opt^ens• , one ol wnicri
Now-
of other subgenera, as a distinct genus.
his
endemic nephrodioides). Although these
{S.
is
adays, pteridoloeists follow Chrislensen's original
u
r
^ rn mountam. s d, o not ih_ arbi or a ilarge number ot species,
^
.
. . c> ' •
placement, giving Stigmatopteris generic sta-
full
they could be considered a secondary regional cen-
tus.
many
Tryon's scheme because they have
ter in
monograph
Christensen^s (191 Dryopteris also
3)
end emic spe
cies.
widened the definition of Stigmatopteris by in-
number
The region with the next highest of
cluding three species what he informally called
in
species Central American, with seven species,
rii-
is
the ^"^ Peltochlaena group," which differed from -/c ,\
j *u
xW^ri-t^xh.
,*^ one which endemic cont,ractu). the
ot \o.
is
true Stigmatopteri.s ,by havi. ng an .ind1 us.ium and1 a
exception of S. contractu, all of the other species
now
nonglandular lamina. This group con-
thick,
is
Andean
extend southward into the Region, giving
Cyclodium, which
sidered part of a separate genus,
the two regions a high floristic affinity.
The
was recently monographed by Smith (1986).
Southeastern Brazil has played a major role in
species of Cyclodium previously placed in Stig-
Stigmatopteris because has
the diversification of
it
matopteris are the excluded taxa section.
listed in
The
six species, of which are endemic. Greater
all
have been important, having three
Antilles also
Geography
which endemic.
species, of are
all
Stigmatopteris entirely neotropical, extend- The Guianan and Mexican centers contain few
is
ing from the Antilles and the state of Veracruz, species and thus have not been important In the
The Mexican Re-
Mexico, southward to southeastern Brazil (Fig. diversification of Srig-ma^op/erts.
1).
The genus absent from the Amazon basin except gion boasts one endemic (S. chimalapensis\ but
is
may
along the western portion adjacent to the Andes. known only from one collection and even-
it is
primarily a montane genus, occurring in wet tually be found outside of the region,
It is
from 200 2,000 m, with the highest
forests to
m
being 2,500 recorded rotundata
elevation for S.
MORPHOLOGY
Panama.
in
The
and endemism Stigmatop- Internal glands the lamina. glands
Species diversity in in
can be conveniently discussed using the re- can usually be seen with a dissecting microscope
teris
centers of high endemism and species rich- and, in oblique light, they appear as tiny ed
gional
bumps They
ness delimited by Tryon (1972) for tropical on the leaf surface (Fig. 3a, c). often
way
American The region of highest dry blackish. Perhaps the best to see the glands
ferns (Fig. 2).
Volume Number 4 Moran 859
78,
1991 Monograph
Stigmatopteris
AMERICA
TROPICAL
map
Flora Neotropica base no
1
2dO 400 —aoC AOO OOOtim
— 1
' I J J
t f " 'T . T r^ 1
TOO 200 300 400 &00 aOO mil»»
Prepared by Hendrik Ryp^efna
fl
O 979 by tha Urivarsttv of Utreclit PijbtJshfid by the Slate University ot Utrecht, the Netherlands Department of Systematic Botany
1
Figure Stigmatopteris.
Distribution of
1.
to look through a hand lens with the lamina guishes Stigmatopteris from other genera of
is all
illuminated from behind. Viewed this way, the glands dryopteroid ferns the presence of uniseriate cilia
is
The
appear as pellucid dots in the lamina (Fig. 3b). that end in a bulbous, often orange or brownish,
i
more some
glands are conspicuous in species than glandular cell (Fig. 4). In other dryopteroid ferns,
in others. Occasionally, they are so large and abun- marginal teeth or processes are formed by the
dant that they are visible to the naked eye, but in projecting end walls of two adjacent cells (Moran,
S. opaca, a species with a thick lamina, the glands 1987, 7) and are never uniseriate or terminated
fig.
The named
are not visible. genus for these glands: by a glandular
is cell.
from the Greek stigmatos, punctate, and pteris^ The scales of Stigmatopteris are more uniform
than those other genera of dryopteroid
fern. in ferns.
Other genera of dryopteroid ferns also have Although the shape differs, species of Stig-
all
internal glands, but these are extremely small and matopteris have scales that are thin, flaccid, and
visible only by thin-sectioning and viewing under tan or light orangish.
a compound microscope (Widen et aL, 1983). The The scales decrease in size from the proximal
become
large glands in Stigmatopteris are probably ho- to distal portions of the lamina until they
2-3
mologous with these smaller ones, but nothing uniseriate or only cells wide at the base (Figs.
is
known about the glands' structure, development, 4, 12b). These highly reduced scales, termed
on
chemical contents, or role in metabolism. **proscales'' or "microscales," are present all
Scales. The scale characteristic that distin- species of the genus, and a dissecting microscope
860 Annals of the
Garden
Missouri Botanical
TROPICAL AMERICA Flora Neotropica base map no
1
O 1979 by tha Univsnity of Ulr«ct)t Publnhad by the Stats University o' Utrecht, the Netherlands Department ot S/itemstic Boleny
and endemism
Figure 2. Distribution of Stigmatopteris species within regional centers of diversity for tropical
American ferns defined by Tryon (1972). Primary centers are indicated with dark shading, secondary centers with
light shading. The number on the left is the total number of species, and the number in parentheses indicates how
many
endemic.
of those species are
needed to see them. Their lateral processes are pinnae with long-acuminate apices that are serrate
is
The symmet-
usually terminated by glandular cells. to the tip. pinnae are equilaterally
Veins. Unlike most dryopteroid ferns, the veins rical, but some of the large species such as S.
may
of end behind the margin in clavate contractu and 5. lechleri have basal pinnae
Sf/g'mafo/)?6?a".s
The
apices which are best seen adaxially (Fig. 23a). All that are elongated basiscopically. texture of
other genera of dryopteroid ferns (sensu stricto) the lamina thin and papyraceous, except for that
is
have the veins ending at or close to the margin, of S. opaca, which thick and slightly charta-
is
and the apices are slender, not thickened. ceous.
Indusia. Stigmatopteris nonindusiate. Three Hairs. All species of Stigmatopteris lack hairs
is
species have an indusiumlike scale associated with except in the adaxial grooves of the rachis and
mm
These 0.05-0.2
the sorus (Figs. 17b, 18a), but these scales are costae. hairs are long, 1- or
They
homologous unbranched, and
certainly not with the large, firm, thick- 2-celled, colorless to reddish.
textured, circular Indusia found in the outgroup are also found in the same position in Cyclodium,
genus Cyclodium. Although independently derived Olfersia, and Polyhotrya and are a synapomorphy
Stigmatopteris^ the lack of an indusium not that holds these three genera and Stigmatopteris
in is
unique because many other dryopteroid ferns also on the same clade within the Dryopteridaceae (see
lack an indusium. evolutionary relationships section). Stigmatopteris
Lamina shape and texture. All species have heterophlebia and S. bulbifera, however, are ex-
Volume Number 4 Moran
78, 861
Monograph
1991 Stigmatopteris
Table 1. Distribution of Stigmatopteris by country. Table 2. The species of Stigmatopteris and their
Endemics are in boldface. distribution by country.
Mexico: 3 species, 1 endemic, chimalapensis, longi- 1. S, brevinervis (Fee) R. C. Moran: southeastern Bra-
caudata, sordida.
zil.
Guatemala: 2 species, no endemics, longicaudata, sordida. 2. S. bulbifera R. C. Moran: Colombia, Ecuador.
Honduras: 1 species, not endemic. longicaudata. 3. S. caudata (Raddi) C. Chr.: southeastern Brazil.
&
Nicaragua: 2 species, no endemics, heterophlebia, sordida. 4. S. chimalapensis Mickel Beitel: southern Mexico.
Costa Rica: 5 species, no endemics, contracta, hetero- 5. S. contracta (Christ) C. Chr.: Costa Rica, Panama.
gemmipara
phlebia, killipiana, longicaudata, sordida. 6. 5. C. Chr.: Jamaica, Haiti.
Panama: 6 species, no endemics, contracta, heterophle- 7. S. hemiptera (Maxon) C. Chr.: Cuba.
bia, killipiana, longicaudata, michaelis, sordida. 8. S. heterocarpa (Fee) Rosenstock: southeastern Bra-
hemiptera.
Cuba: endemic,
species,
1 1 zil.
Jamaica: 2 species, endemic, gemmipara, jamaicensis. 9. S. heterophlebia (Baker) R. C. Moran: Nicaragua,
1
Haiti: species, not endemic, gemmipara. Costa Rica, Panama, Colombia, Ecuador, Peru.
1
Guadeloupe: 1 species, not endemic, rotundata. 10. S. ichthiosma (Sodiro) C. Chr.: western Ecuador.
Dominica: species, not endemic, rotundata. 11. S. jamaicensis (Desv.) Proctor: Jamaica.
1
Martinique: species, not endemic, rotundata. 12. S. killipiana Lellinger: Costa Rica, Panama, Colom-
1
St. Lucia: species, not endemic, rotundata. bia, Ecuador.
1
St. Vincent: species, not endemic, rotundata. 13. S. lechleri (Mett.) C. Chr.: Costa Rica, Venezuela,
1
Grenada: species, not endemic, rotundata. Colombia, Ecuador, Peru.
1
Trinidad: 1 species, not endemic, rotundata. 14. S. longicaudata (Liebm.) C. Chr.: Mexico, Guate-
French Guiana: species, not endemic, rotundata. mala, Honduras, Costa Rica, Panama, French Gui-
1
Guyana: species, not endemic, longicaudata. ana, Venezuela, Colombia, Ecuador, Peru, Bolivia.
1
Panama,
Venezuela: 5 species, 1 endemic, lechleri, longicaudata, 15. S. michaelis (Baker) C. Chr.: Colombia,
nephrodioides, pterorhachis, rotundata. Ecuador.
Colombia: 10 species, no endemics, bulbifera, hetero- 16. S. nephrodioides (Klotzsch) C. Chr.: Venezuela.
opaca
phlebia, ichthiosma, killipiana, lechleri, longicaudata, 17. S. (Baker) C. Chr.: Ecuador, Peru.
michaelis, pterorhachis, pellucidopunctata, sordida. 18. S. pellucidopunctata (C. Chr.) C. Chr.: Colombia,
Ecuador: 10 species, no endemics, bulbifera, heterophle- Ecuador, Peru, Bolivia.
bia, ichthiosma, killipiana, lechleri, longicaudata, mi- 19. S. prionites (Kunze) C. Chr.: southeastern Brazil,
chaelis, opaca, pterorhachis, pellucidopunctata, sordi- 20. S. pterorhachis R. C. Moran: Venezuela, Colombia,
da. 21, S. rotundata (Willd.) C. Chr.: Guadeloupe, Domini-
Peru: 5 species, no endemics, heterophlebia, lechleri, lon- ca, Martinique, St. Lucia, St. Vincent, Grenada, Trin-
gicaudata, opaca, pellucidopunctata. idad, French Guiana, Guyana, Venezuela.
Bolivia: 2 species, no endemics, longicaudata, pellucido- 22. 5. sordida (Maxon) C. Chr.: Mexico, Guatemala,
punctata. Nicaragua, Costa Rica, Panama, Colombia, Ecuador.
Brazil (southeastern): 6 species, 6 endemics, breviner- 23. S. tyucana (Raddi) C. Chr.: southeastern Brazil.
caudata, heterocarpa, prionites, tyucana, 24. Sehnem: southeastern
vis, S. ulei (Christ) Brazil.
ulei.
examined from
Spores. the spores spe-
I 1 1
ceptions to the general condition of hairlessness in cies of Stigmatopteris with the scanning electron
the genus because their axes are covered abaxially microscope (SEM). All of the spores were typical
with capitate-glandular hairs. of those found elsewhere in the Dryopteridaceae,
Sporangia. The sporangia are those de- being monolete, having a smooth exospore, and
like
Moran
scribed by (1987) for Polybotrya, except showing a broadly inflated or narrowly folded peri-
The
that the stalks lack a hairlike paraphysis. spore (Fig. 5). irmer layer of the perispore is
A
Petiole anatomy. cross section of the pet- thin and usually adheres to the smooth exospore;
shows that the vascular bundles are arranged the outer layer thicker and compact,
iole is
mushroomlike The on and between
in a outline, with the base of the perisporal surface the folds
mushroom oriented adaxially (Fig. 12a). This was nearly uniform in species examined (Fig.
is all 1 1
The
typical of dryopteroid ferns (Moran, 1986, 1987; surface relatively unornamented, being
5). is
pers, obs.). Aerophores run down the sides of the either smooth (Fig. 5f) or beset with low, diffuse
petiole and are evident in cross section by a gap ridges (Fig. 5c, d). Because of the uniformity, the
in the thickened peripheral coUenchyma of the spores do not provide helpful characters in the
petiole. The petiole bases do not persist or accu- subgeneric taxonomy of the genus,
Ornamentation
mulate starch after the rest of the leaf has withered. of the perispore differs in Stig-
In other words, Stigmatopteris lacks trophopods. matopteris from that of the most closely related
862 Annals the
of
Garden
Missouri Botanical
FuiURE 3. Internal punctate glands in the lamina of Stigmatopteris (a-c) and rachis-costa juncture showing
—
mm.
White (0llgaard 35866,
adaxial grooves pubescent with short hairs (d). hars indicate 1 a, b. 5. bulbifera et al.
—
—
AAU). contracta (Moran 3330, MO). heterocarpa {Ilaerchen 106,
S, d. S. B).
c.
Volume Number 4 Moran
863
78,
1991 Monograph
Stigmatopteris
Figure The shown from from
4. Scales of Stigmatopteris. scale in b is a petiole base; all others are the abaxial
surface of the lamina and show the transition from uniseriate scales (proscales or microscales) to multiseriate ones.
—
All are the same magnification as shown in d, except b and the far right scale in a. a. S. pellucidopunctata (Moran
— —
—
3624, MO). b. S. ichthiosma {Moran 3545, MO). c. S. sordida (Gomez 18641, CR). d. S. lechleri (Moran
—
MO).—
3582, MO). opaca (Moran 3576, sordida (Moran 3315, MO).
e. S. f. S.
864 Annals of the
Garden
Missouri Botanical
Volume Number 4 Moran 865
78,
Monograph
1991 Stigmatopteris
CYCLODIUM STIGMATOPTERIS OLFERSIA POLYBOTRYA
dryopteroid genera, Polybotrya, Cyclodium,
i.e.,
and These genera have more ornamen-
Olfersia.
tation on the perispore. For example, the spores
of Polybotrya are typically densely echinate (Mo-
&
ran, 1987; Tryon Tryon, 1982); those of Cy-
clodium are usually finely papillate but can be
&
variously ornamented (Smith, 1986; Tryon
and
Tryon, 1982); those of Olfersia are sparsely
&
Tryon
echinate (Moran, 1986; Tryon, 1982).
Although these types of spore ornamentation are
premature determine which type
distinct, to
it is
Figure 6. Cladogram of Cyclodium, Stigmatopteris^
and which Such
ancestral derived. a deter-
is is
Olfersia, and Polybotrya. The numbered black bars rep-
mination must await a comprehensive comparative
The
resent the derived character states given in Table 4.
ornamentation Dryop-
study of perisporal in the
open bar below Olfersia represents a character state re-
teridaceae. Therefore, have not used spore or- versal. See Moran (1986) for the additional apomorphies
I
namentation cladogram Stigmatopteris and synapomorphies pertaining to Olfersia and Polybo-
in the for
trya.
and genera
related (Fig.
6).
Chromosome Number
hairs small catenate, several-celled hairs
tis (i.e.,
Walker (1966), who has made the only chro- with the color aggregated in the cross walls), and
=
mosome count for Stigmatopteris, found that n having sulcate, decurrent axial grooves.
41 in S. jamaicensis (reported as S. nothochlaena. Generic relationships. Stigmatopteris most
a synonym). This number relates the genus to the closely resembles Cyclodium (sensu Smith, 1986),
dryopteroid another dryopteroid genus, as shown by similarities
ferns.
Walker also noted that meiosis occurs very early in leaf cutting and venation and by the laminae,
Stigmatopteris during the crozier stage of leaf which are hairless except for the grooves of the
in
development. Unfortunately, most workers rachis and costae. In fact, the two genera are so
(in-
some
eluding myself) have collected samples for chro- similar in these respects that pteridologists
&
mosome Christensen, 1913, 1920; Tryon Tryon,
counts past the crozier stage, at a stage (e.g.,
more typical for ferns in general, when the 1982) have regarded them as one. They are, how-
i.e.,
The
are small and white, but on expanded leaves ever, best kept as distinct. two genera exhibit
sori
beyond the crozier stage. This probably explains 11 morphological dififerences (Table and this
3), is
why only one count exists for the genus. more than that which occurs between other pairs
of dryopteroid genera, such as Arachniodes and
Dryopteris, Arachniodes and Maxonia, or Cyr-
Evolutionary Relationships
tomium and Polystichum. If there to be con-
is
Family relationships. That Stigmatopteris sistency in recognizing dryopteroid genera, at least
is
member number
a of the Dryopteridaceae (sensu stricto of as regards the of characters that separate
Holttum, 1984) clearly shown by several char- them, then Cyclodium and Stigmatopteris should
is
acteristics which shares with genera of that be maintained as distinct.
it all
=
family: x 41, more than three vascular bundles In the Dryopteridaceae, Stigmatopteris and Cy-
omega
and arranged an shape clodium form Polybotrya and
in the petiole these in a clade with Olfersia
when
viewed in cross section (Fig. 2a), and grooves (Fig. 6, Table 4). This clade defined by the
1 is
on the adaxial surface of the axes that are decur- presence of hairs in the adaxial grooves of the axes
The Dryop-
rent into those of the next lower order. (Fig. 3d). This character state considered syn-
is
teridaceae are closely related to the Tectariaceae apomorphic because other dryopteroid ferns
all
(sensu stricto of Holttum, 1984), but Stigmatop- (sensu stricto) are hairless in the adaxial grooves
teris differs from the latter family by lacking Cteni- of the axes. This character state would be weak
if
—
Figure 5. Spores oi Stigmatopteris.^ a, c. S. longicaudata {Mickel 6392^ NY). b. S. lechleri {von Sneidern
—
— —
5046, US). tyucana (Brade 5892, NY). opaca (0llgaard 35859, AAU). sordida {Moran
d. S. e. S. et at. f. S.
—
3315, MO).- rotundata {Cremers 9140, UC). caudata {Webb NY). Spores c-h the same scale
5. h. S. s.n.,
g.
as
in b.
866 Annals of the
Garden
Missouri Botanical
Table Comparison Stigmatopteris and Cjclodium.
of
3.
Similarities
and
Hairs of the lamina: lacking (except in grooves of the rachis costae).
mm
Hairs grooves of the rachis and costae: 0.05-0.2 long, usually blunt-tipped, colorless, one- to few-celled.
in
Lamina cutting: similar in many species, cf. S. rotundata and C. guianense, or S. opaca and C. meniscioides.
Venation: similar in free-veined species such as S, nephrodioides and C. guianense, and in anastomosing-veined
species such as S. heterophlehia and C. meniscioides.
Differences
Cyclodium
Character Stigmatopteris
present
Indusium; absent
Lamina papyraceous chartaceous or coriaceous
texture:
anadromous catadromous anadromous throughout
Pinnule arrangement basally,
distally
margm
behind margin
Position of vein ending: at
Shape of vein clavate slender
tips:
Internal laminar glands: present absent
common
Scale frequency on rare or lacking
*
brown orangish dark brown
Scale color: light or
thick
Scale texture: thin
composed upturned margins
Marginal teeth of scales: uniseriate of
from two adjacent walls
cell
Gland at the apex of scale teeth: present absent
Andean Guianan
Geography: primarily primarily
nothing else correlated with could have evolved of Polybotrya subg. Polybotrya, such as P. os-
it; it
much
more than once, and so the clade would be miindacea and P. cyatheoides, have longer
if
polyphyletic. The hypothesis that character hairs the grooves (Moran, 1987, 49c) than
this in fig.
evolved only once supported by the fact that the those found in the grooves of Cyclodium and 5^/^--
is
type of pubescence the same the genera; matopteris. These longer hairs, however, are oth-
in
is all
mm
the hairs are 0.05-0.2 long, usually blunt- erwise similar to the smaller ones and are probably
tipped, colorless, and one- to few-celled (Smith, derived from them.
1986; Moran, 1987, lOb-e, Oljersia The clade with Stigmatopteris, Olfersia, and
fig. h).
g,
cervina, the only species of Oljersia, an excep- Polybotrya defined only by one synapomorphy:
is is
tion, having glabrous grooves. Because Olfersia loss of the indusium (Fig. 6, Table 4). This synapo-
is
the sister genus to Polybotrya (Moran, 1986), morphy weak and therefore considered tentative
its is
lack of hairs the grooves best Interpreted as because no others correlate with Furthermore,
In
is it.
Some
a character stale reversal (Fig. species the number of times the indusium has been lost in
6).
Table 4. Characters and character states used in the cladistic analysis of Stigmatopteris, Cyclodium, Olfersia,
and Polybotyra Character were determined by comparison other genera the Dryopteridace-
(Fig. state polarities to in
6).
such as Arachniodes, Dryopteris, and Polystichum.
ae,
Character Ancestral condition Derived condition
Pubescence adaxlal grooves of the axes absent present
in
1.
Indusium present absent
2.
Large glands the lamina absent present
3. in
composed upturned margins
4. Marginal teeth or processes of the scales of the uniseriate
from two adjacent
cells
5. Glandular cell at apex of scale teeth or absent present
processes
margm
behind margin
Vein termination
at
6.
Shape of vein slender clavate
7. tip
membranaceous
8. Texture of lamina papyraceous or chartaceous
