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A Classification for Extant Ferns



We present a revised classification for extant ferns, with emphasis on ordinal and familial ranks, and a synopsis of included genera. Our classification reflects recently published phylogenetic hypotheses based on both morphological and molecular data. Within our new classification, we recognize four monophyletic classes, 11 monophyletic orders, and 37 families, 32 of which are strongly supported as monophyletic. One new family, Cibotiaceae Korall, is described. The phylogenetic affinities of a few genera in the order Polypodiales are unclear and their familial placements are therefore tentative. Alphabetical lists of accepted genera (including common synonyms), families, orders, and taxa of higher rank are provided.
Recent phylogenetic studies have revealed a basal
dichotomy within vascular plants, separating the lyco-
phytes (less than 1% of extant vascular plants) from the
euphyllophytes (Fig. 1; Raubeson & Jansen, 1992; Ken-
rick & Crane, 1997; Pryer & al., 2001a, 2004a, b). Liv-
ing euphyllophytes, in turn, comprise two major clades:
the spermatophytes (seed plants), which are in excess of
260,000 species (Thorne, 2002; Scotland & Wortley,
2003), and the monilophytes (ferns, sensu Pryer & al.,
2004b), with about 9,000 species, including horsetails,
whisk ferns, and all eusporangiate and leptosporangiate
ferns. Plants that are included in the lycophyte and fern
clades are all spore-bearing or “seed-free”, and because
of this common feature their members have been lumped
together historically under various terms, such as “pteri-
dophytes” and “ferns and fern allies”—paraphyletic
assemblages of plants. The focus of this reclassification
is exclusively on ferns (Division Tracheophyta, Sub-
division Euphyllophytina, Infradivision Moniliformop-
ses, of Kenrick & Crane, 1997), characterized by lateral
root origin in the endodermis, usually mesarch pro-
toxylem in shoots, a pseudoendospore, plasmodial tape-
tum, and sperm cells with 30–1000 flagellae (Renzaglia
& al., 2000; Schneider & al., 2002a).
Increasingly robust phylogenetic hypotheses for
ferns (Hasebe & al., 1994, 1995; Manhart, 1994, 1995;
Pryer & al., 1995, 2001a, 2004b; Kranz & Huss, 1996;
Pahnke & al., 1996; Wolf, 1997; Wolf & al., 1998;
Beckert & al., 1999; Vangerow & al., 1999; Sano & al.,
2000a; Schneider & al., 2004c; Wikström & Pryer, 2005;
Tsutsumi & Kato, 2006; Schuettpelz & al., in press), uti-
Smith & al. • Fern classification55 (3) • August 2006: 705–731
Fig. 1. Consensus phylogeny depicting relationships of
major vascular plant lineages. Topology summarizes the
results of previously published phylogenetic studies
(e.g., Raubeson & Jansen, 1992; Kenrick & Crane, 1997;
Renzaglia & al., 2000; Pryer & al., 2001a, see main text for
others). Resolved nodes have received bootstrap support
A classification for extant ferns
Alan R. Smith1, Kathleen M. Pryer2, Eric Schuettpelz2, Petra Korall2,3, Harald Schneider4&
Paul G. Wolf5
1University Herbarium, 1001 Valley Life Sciences Building #2465, University of California, Berkeley,
California 94720-2465, U.S.A. (author for correspondence).
2Department of Biology, Duke University, Durham, North Carolina 27708-0338, U.S.A.
3Department of Phanerogamic Botany, Swedish Museum of Natural History, Box 50007, SE-104 05 Stock-
holm, Sweden.
4Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Abteilung Systematische Botanik, Georg-August-
Universität, Untere Karspüle 2, 37073 Göttingen, Germany.
5Department of Biology, Utah State University, Logan, Utah 84322-5305, U.S.A.
We present a revised classification for extant ferns, with emphasis on ordinal and familial ranks, and a synop-
sis of included genera. Our classification reflects recently published phylogenetic hypotheses based on both
morphological and molecular data. Within our new classification, we recognize four monophyletic classes, 11
monophyletic orders, and 37 families, 32 of which are strongly supported as monophyletic. One new family,
Cibotiaceae Korall, is described. The phylogenetic affinities of a few genera in the order Polypodiales are
unclear and their familial placements are therefore tentative. Alphabetical lists of accepted genera (including
common synonyms), families, orders, and taxa of higher rank are provided.
classification, Cibotiaceae, ferns, monilophytes, monophyletic.
lizing data from morphology, seven chloroplast markers
(rbcL, atpA, atpB, accD, rps4, 16S rDNA, ITS), one nu-
clear gene (18S rDNA), and three mitochondrial genes
(atp1, nad2, nad5) prompt us to reevaluate the classifi-
cation of these vascular plants. Multiple-gene phyloge-
netic analyses, e.g., studies by Wolf (1996), Wolf & al.
(1998), Pryer & al. (2001a, 2004b), Schneider & al.
(2004c), Wikström & Pryer (2005), and Schuettpelz & al.
(in press), have given rise to growing conviction in both
the composition and relationships of taxa at familial and
ordinal ranks. Five recent morphological analyses of
relationships, by Pryer & al. (1995, 2001a), Schneider
(1996a), Stevenson & Loconte (1996), and Schneider &
al. (in prep.) have increased support for the molecular-
based consensus topology. For eusporangiate and basal
leptosporangiate ferns, evidence is now sufficient to al-
low us to circumscribe confidently most clades and as-
sign ranks. However, for some more derived leptospo-
rangiate ferns, the phylogenetic evidence is still some-
what equivocal. Consequently, declaration of phyloge-
netic positions for some taxa and the assignment of ranks
(which we consider subjective and secondary), in a clas-
sical “Linnaean-style” hierarchy, are tentative. In this pa-
per, we present a revised view of the classification of ex-
tant ferns, taking into account all relevant evidence. We
focus our classification at the ranks of class, order, and
family, believing that the information at hand is most
appropriate for resolution and understanding of relation-
ships at these levels. Within most families, and especial-
ly at the generic level, there is still insufficient evidence
to attempt many classificatory decisions.
In the classification proposed herein, we account for,
and place in a revised taxonomic framework, all names at
family and ordinal rank utilized in previous major classifi-
cations directed at ferns, particularly those that have been
proposed in the last eighty years: Bower (1926), Christen-
sen (1938), Ching (1940, 1978), Dickason (1946), Cope-
land (1947), Holttum (1947, 1949, 1973), Pichi Sermolli
(1958, 1977), Mehra (1961), Wagner (1969), Nayar
(1970), Tagawa & Iwatsuki (1972), Mickel (1974), Try-
on & Tryon (1982), Kramer (in Kubitzki, 1990), Hennip-
man (1996), and Stevenson & Loconte (1996). We also
consider various herbarium schemes in common use,
such as the one by Crabbe & al. (1975). Many of these
classifications, as well as others, have been reviewed and
reproduced by Pichi Sermolli (1973), who also summa-
rized detailed information on family names of ferns
(1970, 1981, 1982, 1986). Smith (1995) provided a re-
cent summary and discussion of these classifications.
Classifications serve many purposes, among them to
provide a genealogical framework in which to identify
plants, organize herbaria, retrieve information, and to
conduct many kinds of studies (e.g., evolutionary, mor-
phological, and physiological). Multi-use classifications
serve us best if we name only those clades that are read-
ily recognizable and characterized by morphological
synapomorphic characters, at least at family and higher
ranks. However, more traditional (morphology-based)
and practical classifications are sometimes incompatible
with the results and classifications implied by phyloge-
netic studies, especially when the principle of monophy-
ly is used as a grouping criterion (recognizing clades, and
not paraphyletic grades; APG II, 2003). When a tradi-
tionally recognized family nests within another, complex
classificatory choices ensue: (1) recognition of para-
phyletic families (Brummitt, 1996, 1997; Lidén & al.,
1997; Moore, 1998; Diggs & Lipscomb, 2002; Grant,
2003); (2) dismemberment of a recognized family into
smaller families (e.g., the disintegration of classical
Scrophulariaceae; Olmstead & al., 2001); or (3) integra-
tion of the traditional family that causes the paraphyly
into the “progenitor” family. The first choice, preferred
by some, leads to recognition of unnatural (non-mono-
phyletic) groups, which in our opinion often retards or
obscures investigation into interesting biological, phyto-
geographic, and evolutionary questions. The second so-
lution supposes that we have morphological synapomor-
phies for nodes that lead to all of the segregate families,
and this is not often the case, although one hopes that
eventually we shall find these synapomorphies. Until
then, it may be nearly impossible to define some segre-
gate families in such a way that they would be both
keyable and circumscribable. The third solution is a “fast
fix” to the problem, but expediency often demands that
at least some of the intrafamilial subclades also be rec-
ognized taxonomically, either at a lower rank (e.g., sub-
family), or with an unranked informal name (e.g., “gram-
mitid ferns”) until further decisions on rank can be made.
This third option seems to us the most practicable and
practical solution toward a “first-pass” revision of fern
classification. As more data are gathered and future phy-
logenetic analyses provide better resolved and better sup-
ported topologies, one expects further insight into identi-
fying synapomorphies for segregate taxa, enabling even-
tual movement toward the second option of recognizing
segregate monophyletic families.
The nested linearity of a ranked Linnaean classifica-
tion and the availability of a limited number of univer-
sally accepted ranks facilitate the organization and ar-
rangement of taxa for cataloging purposes, such as the
efficient arrangement of specimens in herbaria or the
organization and retrieval of regional biodiversity from
checklists, floras, and other taxonomic products. Ranked
classifications often perform poorly, however, when
called upon to present explicit statements of clade mem-
bership and relationship. Such classifications also can
mislead one into making specious comparisons, e.g., at-
tempting to describe the extent of biodiversity in terms of
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
numbers of taxa of a particular arbitrarily designated
rank (comparing apples with oranges). Such problems
notwithstanding, it is nevertheless highly likely that
ranked Linnaean classifications will continue to co-exist
with phylogenetic classifications for the foreseeable
In this paper we combine the principle of monophy-
ly with a desire to maintain well-established names to
update ordinal and familial ranks within ferns so that
they are better reconciled with our current best estimates
of phylogenetic relationships. However, we attempt to
avoid recognizing monogeneric families within the more
derived leptosporangiate ferns. To paraphrase Crisp & al.
(1999), who addressed the matter of monotypic genera,
the most compelling case for recognizing a monotypic
family is when such a family is characterized by one or
more autapomorphies and is the sister group to a clade of
families, all of the members of which lack these apomor-
phies. The alternative would be to treat the whole clade
(including the autapomorphic family) as a single family,
which results in sinking some well-circumscribed fami-
lies into synonymy. At this time, we prefer to utilize a
minimum number of ranks to categorize only the most
well supported splits in the phylogeny.
In some respects, our new classification differs in
relatively minor ways in the circumscription of orders,
families, and hierarchical structure from most other re-
cent fern classifications (e.g., those of Tryon & Tryon,
1982; Kramer, in Kubitzki, 1990; and Stevenson &
Loconte, 1996), but there are significant departures. Only
one previous fern classification has employed cladistic
methodology in a rigorous way; Stevenson & Loconte
(1996) superimposed on their tree a hierarchical classifi-
cation (albeit somewhat abbreviated), but the phylogeny
that they generated was based exclusively on morpho-
logical data and differs radically from the most up-to-
date phylogenetic hypothesis. Our classification, in con-
trast, is based on consensus of a variety of morphologi-
cal and molecular studies.
Fern names above the rank of genus used in this clas-
sification (Appendices 1–4) have been obtained, for the
most part, from the web site of James Reveal, University
of Maryland (
PBIO/fam/hightaxaindex.html) and from Hoogland &
Reveal (2005). Most names at family rank are also listed
and discussed by Pichi Sermolli (1970, 1982) and sum-
marized in reports by the Subcommittee for Family
Names of Pteridophyta (Pichi Sermolli, 1981, 1986); this
list was further emended and updated by Pichi Sermolli
(1993). To the best of our knowledge (and unless other-
wise indicated), the family names in Appendix 3 have all
been published in accord with the International Code of
Botanical Nomenclature (Greuter & al., 2000) and thus
satisfy the relevant Articles in the Code with regard to
publication and priority of family names (Art. 10.6, Art.
11.1–11.3; Art. 18; Art. 41.1) and names of ranks above
family (Art. 16, 17). Although the principle of priority is
not mandatory for names of taxa above the rank of fam-
ily (Art. 11.9), authors are advised to follow this princi-
ple (Rec. 16B.1). Of the orders we recognize below, all
have priority except Salviniales, which is later than the
much less commonly applied name Pilulariales.
The rules of nomenclature applicable to supra-ordi-
nal names are the same as those for ordinal names, dis-
cussed above. We have made no attempt to scrutinize the
literature for names above the rank of order, but many of
these names are included in lists available from the
Reveal website (and so are included in Appendix 1) and
in Hoogland & Reveal (2005); still others can be gleaned
from Kenrick & Crane (1997, see, especially, their Table
7.2, pp. 231–233; however, many of these are not valid-
ly published) and from Pichi Sermolli (1959). Names at
ordinal and subordinal ranks, also mostly from the Re-
veal website and from Hoogland & Reveal (2005), are
given in Appendix 2, names at familial rank are given in
Appendix 3. Citations for all names are given in Ap-
pendix 4. We also present an index to commonly accept-
ed genera with family assignments proposed here
(Appendix 5).
In the classification that follows, for each family, we
give common names (often derived from the scientific
names) for the clades (if there is one that has common
usage), heterotypic synonyms, approximate numbers of
genera and species, names of constituent genera, refer-
ences to relevant phylogenetic literature, and discussion
of unresolved problems, where appropriate. DNA se-
quence data are now available for all families recognized
herein, and for most genera of ferns. A superscript num-
ber one (
) denotes those genera for which DNA se-
quence data are not available; nonetheless, taxonomic
placement for most of these is relatively certain, based on
morphological evidence. Lack of a superscript indicates
that some molecular evidence (either published or
unpublished) has been available to us for consideration
and we are relatively confident in the placement of the
genus. The classification presented below is based on the
consensus relationships depicted in Fig. 2, which are
derived from, and guided by, recent and ongoing phylo-
genetic studies (e.g., Hasebe & al., 1995; Pryer & al.,
2004b; Schneider & al., 2004d; Korall & al., 2006;
Schuettpelz & al., 2006, unpubl. data).
In this reclassification, we treat all classes, orders,
and families of extant ferns, which constitute a mono-
Smith & al. • Fern classification55 (3) • August 2006: 705–731
phyletic group, sometimes referred to as Infradivision
Moniliformopses (Kenrick & Crane, 1997), or monilo-
phytes (Donoghue in Judd & al., 2002; Pryer & al.,
2001a, 2004a, b). However, “Infradivision” is not a rec-
ognized rank in the International Code of Botanical No-
menclature (Greuter & al., 2000); moreover, the name
“Moniliformopses” was never validly published, lacking
a Latin diagnosis or description, or a reference to one.
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
Fig. 2. Consensus phylogeny depicting fern relationships upon which our classification is based. Topology summar-
izes the results of previously published and ongoing phylogenetic studies (e.g., Hasebe & al., 1995; Pryer & al., 2004b;
Schneider & al., 2004b; Korall & al., 2006; Schuettpelz & al., in press; Schuettpelz, unpubl.; see main text for others).
Resolved nodes have received bootstrap support 70 except those drawn with dotted lines. Family, order, and class
names that correspond to our classification are indicated to the right. Common names for some larger clades are indi-
cated to the left.
Ophioglossaceae Psilotopsida
Core leptosporangiates
Eupolypods II
Eupolypods I
Because validly published names at an appropriate rank
are not available for ferns (as here defined), euphyllo-
phytes (ferns + seed plants), or even vascular plants (eu-
phyllophytes + lycophytes), and because all available
names at higher ranks have been used in a sense we think
would be confusing, we avoid placing ferns in any rank
above class. Classification may be better served by adop-
ting “floating”, rankless names for clades above the rank
of class.
Within ferns, we recognize four classes (Psilotop-
sida; Equisetopsida; Marattiopsida; Polypodiopsida), 11
orders, and 37 families.
1. Family Ophioglossaceae. — Ophioglosso-
ids; incl. Botrychiaceae, Helminthostachyaceae.
Four genera: Botrychium (grapeferns; moonworts), Hel-
minthostachys, Mankyua
, Ophioglossum (adder
tongues). Botrychium (incl. Botrychium s.s., Sceptridi-
um, Botrypus, and Japanobotrychium) and Ophioglos-
sum (incl. Cheiroglossa, Ophioderma) are sometimes di-
vided more finely (Kato, 1987; Hauk & al., 2003). Ca. 80
spp.; monophyletic (Hasebe & al., 1995; Hauk, 1995;
Pryer & al., 2001a, 2004b; Hauk & al. 2003). Mankyua,
from Cheju Island, Korea, has recently been described,
but no molecular data are available (Sun & al., 2001).
Species mostly terrestrial (a few epiphytic), temperate
and boreal, but a few pantropical. Characters: vernation
nodding (not circinate); rhizomes and petioles fleshy;
root hairs lacking; aerophores absent (Davies, 1991); fer-
tile leaves each with a single sporophore arising at the
base of, or along, the trophophore stalk, or at the base of
the trophophore blade (several sporophores per blade in
Cheiroglossa); sporangia large, with walls two cells
thick, lacking an annulus; spores globose-tetrahedral, tri-
lete, many (> 1000) per sporangium; gametophytes sub-
terranean, non-photosynthetic, mycorrhizal; x = 45 (46).
2. Family Psilotaceae. — Whisk ferns; incl.
Two genera (Psilotum, Tmesipte-
ris), ca. 12 total spp. (2 in Psilotum); monophyletic (Ha-
sebe & al., 1995; Pryer & al., 2001a, 2004). Characters:
roots absent; stems bearing reduced, unveined or single-
veined euphylls; sporangia large, with walls two cells
thick, lacking an annulus; two or three sporangia fused to
form a synangium, seemingly borne on the adaxial side
of a forked leaf; spores reniform, monolete, many (>
1000) per sporangium; gametophytes subterranean (Psi-
lotum), non-photosynthetic, mycorrhizal; x = 52.
3. Family Equisetaceae. — Horsetails.
A single
genus (Equisetum), 15 spp. usually placed in two well-
marked subgenera, subg. Equisetum and subg. Hippo-
chaete; monophyletic (Pryer & al., 2001a, 2004b; Des
Marais & al., 2003; Guillon, 2004). The spermatozoids
of Equisetum share several important features with other
ferns that support their inclusion in this clade (Renzaglia
& al., 2000). Kato (1983) adduced additional morpho-
logical characters, including root characters, supporting a
relationship between horsetails and ferns. Characters:
stems whorled, lacunate; leaves whorled, connate; spo-
rangia with helical secondary wall thickenings (Bate-
man, 1991), borne on peltate sporangiophores that col-
lectively comprise strobili; sporangia large, lacking an
annulus, many (> 1000) per sporangium; spores green,
with circular aperture and four paddle-like, coiled ela-
ters; gametophytes green, surficial; x = 108.
4. Family Marattiaceae. — Marattioids; incl.
Angiopteridaceae, Christenseniaceae, Danaeaceae,
Four genera: Angiopteris, Christen-
senia, Danaea, Marattia; Archangiopteris has been rec-
ognized by some (e.g., Pichi Sermolli, 1977) but appears
to nest within Angiopteris (Murdock, 2005); Danaea is
sister to the other three genera (Pryer & al., 2001a,
2004b; Murdock, 2005) and represents a neotropical
radiation (Christenhusz & al., unpubl.); Angiopteris and
Christensenia are restricted to eastern and southeastern
Asia, Australasia, and Polynesia, while Marattia is pan-
tropical. Ca. 150 spp., but monographic revision is need-
ed at the species level in several genera; monophyletic
(Hill & Camus, 1986; Pryer & al., 2001a, 2004b;
Murdock, 2005). We see no advantage or good reason for
recognizing several of the constituent genera as mono-
generic families, as done by Pichi Sermolli (1977). The
name Danaeaceae has been found to predate
Marattiaceae (Appendix 4); however, Marattiaceae is
being proposed for conservation by Murdock & al.
(subm.), and we maintain its usage in the usual broad
Terrestrial (rarely epipetric) pantropical, fossils be-
ginning in Carboniferous (Collinson, 1996). Characters:
roots large, fleshy, with polyarch xylem; root hairs sep-
tate; roots, stems, and leaves with mucilage canals; rhi-
zomes fleshy, short, upright or creeping, with a polycy-
Smith & al. • Fern classification55 (3) • August 2006: 705–731
clic dictyostele; vernation circinate; leaves large, fleshy,
1–3-pinnate (rarely simple in Danaea, or 3–5-foliate in
Christensenia) with enlarged, fleshy, starchy stipules at
the base and swollen pulvinae along petioles and rachis-
es (and sometimes other axes); petiole and stem xylem
polycyclic; stems and blades bearing scales; pneuma-
thodes (lenticels) scattered all around petioles and/or ra-
chises; sporangia free or in round or elongate synangia
(fused sporangia), lacking an annulus, enclosing 1000–
7000 spores; spores usually bilateral or ellipsoid, mono-
lete; gametophytes green, surficial; x = 40 (39).
5. Family Osmundaceae. —
Three genera: Lep-
topteris, Osmunda, Todea. Ca. 20 spp.; monophyletic
(Hasebe & al., 1995; Yatabe & al., 1999; Pryer & al.,
2001a, 2004b). Fossils from Permian; temperate and
tropical. Characters: stem anatomy distinctive, an ecto-
phloic siphonostele, with a ring of discrete xylem
strands, these often conduplicate or twice conduplicate in
cross-section; stipules at bases of petioles; leaves dimor-
phic or with fertile portions dissimilar to sterile; sporan-
gia large, with 128–512 spores, opening by an apical slit,
annulus lateral; spores green,subglobose, trilete; gameto-
phytes large, green, cordate, surficial; x = 22.
6. Family Hymenophyllaceae. — Filmy ferns;
incl. Trichomanaceae.
Nine genera (Ebihara & al.,
2006), two major clades (Pryer & al., 2001b), “trichoma-
noid” and “hymenophylloid”, roughly corresponding to
the classical genera Trichomanes s.l. and Hymenophyl-
lum s.l. Ca. 600 spp.; monophyletic (Hasebe & al., 1995;
Dubuisson, 1996, 1997; Pryer & al., 2001b, 2004b; Ebi-
hara & al., 2002, in press; Dubuisson & al., 2003; Hen-
nequin & al., 2003). Several segregate and monotypic
genera are nested within Hymenophyllum s.l.: Cardio-
manes, Hymenoglossum, Rosenstockia, and Serpyllopsis
(Ebihara & al., 2002, in press; Hennequin & al., 2003).
Several other classically defined hymenophylloid genera
(subgenera) may not be monophyletic, e.g., Mecodium
and Sphaerocionium (Hennequin & al., 2003; Ebihara &
al., 2006). Microtrichomanes appears to be polyphyletic
(Ebihara & al., 2004). Trichomanes s.l. comprises eight
monophyletic groups that are regarded here as genera:
Abrodictyum s.l., Callistopteris, Cephalomanes, Crepi-
domanes, Didymoglossum, Polyphlebium, Trichomanes
s.s., and Vandenboschia; several of these have been sub-
divided into putatively monophyletic subgenera and sec-
tions (Ebihara & al., in press). Terrestrial and epiphytic;
pantropical and south-temperate, but gametophytes sur-
vive in north-temperate regions as far north as Alaska.
Characters: rhizomes slender, creeping, wiry, or some-
times erect and stouter, protostelic; vernation circinate;
blades one cell thick between veins (a few exceptions);
stomata lacking; cuticles lacking or highly reduced;
scales usually lacking on blades, indument sometimes of
hairs; sori marginal, indusia conical (campanulate), tubu-
lar, or clam-shaped (bivalvate), with receptacles (at least
in trichomanoid genera) usually elongate, protruding
from the involucres; sporangia maturing graduately in
basipetal fashion, each with an uninterrupted, oblique
annulus; spores green, globose, trilete; gametophytes fil-
amentous or ribbon-like, often reproducing by fragmen-
tation or production of gemmae; x = 11, 12, 18, 28, 32, 33,
34, 36, and perhaps others.
Dipteridales, Matoniales, Stromatopteridales.
nophyletic (Pryer & al., 2004b; Schuettpelz & al., 2006).
Characters: root steles with 3–5 protoxylem poles
(Schneider, 1996a); antheridia with 6–12 narrow, twisted
or curved cells in walls.
7. Family Gleicheniaceae. — Gleichenioids,
forking ferns; incl. Dicranopteridaceae, Stromato-
Six genera (Dicranopteris, Diplopterygi-
um, Gleichenella, Gleichenia, Sticherus, Stromatopteris),
ca. 125 spp.; monophyletic (Hasebe & al., 1995; Pryer &
al., 1995, 2001a, 2004b). Hennipman (1996) also sugges-
ted inclusion of the next two families in Gleicheniaceae;
however, these are recognized here as distinct based on
their significant morphological disparity. Fossil record
beginning in Cretaceous (Jurassic and older fossils may
belong to the Gleicheniales or represent ancestors of ex-
tant Gleicheniaceae); pantropical. Characters: rhizomes
with a “vitalized” protostele, or rarely solenostele; leaves
indeterminate, blades pseudodichotomously forked (ex-
cept Stromatopteris); veins free; sori abaxial, not mar-
ginal, with 5–15 sporangia, each with a transverse-
oblique annulus, exindusiate, round, with 128–800
spores; sporangia maturing simultaneously within sori;
spores globose-tetrahedral or bilateral; gametophytes
green, surficial, with club-shaped hairs; x = 22, 34, 39,
43, 56.
8. Family Dipteridaceae. — Including Cheiro-
Two genera, Cheiropleuria and Dipteris,
from India, southeast Asia, eastern and southern China,
central and southern Japan, and Malesia, to Melanesia
and western Polynesia (Samoa), ca. 11 spp.; mono-
phyletic (Kato & al., 2001; Pryer & al. 2004b). Fossil
record beginning in upper Triassic. Characters: stems
long-creeping, solenostelic or protostelic, covered with
bristles or articulate hairs; petioles with a single vascular
bundle proximally and polystelic distally; blades (sterile
ones, at least) cleft into two or often more subequal parts;
veins highly reticulate, with included veinlets; sori exin-
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
dusiate, discrete, compital (served by many veins), scat-
tered over the surface, or leaves dimorphic and the fertile
ones covered with sporangia; sporangia maturing simul-
taneously or maturation mixed, with a 4-seriate stalk;
annuli almost vertical or slightly oblique; spores ellip-
soid and monolete, or tetrahedral and trilete, 64 or 128
per sporangium; gametophytes cordate-thalloid; x = 33.
Dipteris differs from Cheiropleuria primarily in having
bilateral, monolete spores (tetrahedral and trilete in
Cheiropleuria) and monomorphic leaves with discrete
sori (sporangia acrostichoid in Cheiropleuria).
9. Family Matoniaceae. — Matonioids.
genera (Matonia, Phanerosorus), each with two spp.;
monophyletic, sister to Dipteridaceae (Kato & Setogu-
chi, 1998; Pryer & al., 2004b; Schuettpelz & al., 2006).
Malesia-Pacific Basin; fossil record beginning in mid-
Mesozoic. Characters: stems solenostelic with at least
two concentric vascular cylinders (polycyclic) and a cen-
tral vascular bundle; blades flabellate (Matonia), un-
evenly dichotomously branched or with dichotomous
pinnae; veins free or slightly anastomosing around sori;
sori with peltate indusia; sporangia maturing simultane-
ously, with very short stalks and oblique annuli; spores
globose-tetrahedral, trilete; gametophytes green, thal-
loid, with ruffled margins; antheridia large, many-celled;
x = 26 (Matonia), 25 (Phanerosorus).
(Hasebe & al., 1995; Pryer & al., 2001a, 2004b; Skog &
al., 2002; Wikström & al., 2002). Fossil record beginning
in the Jurassic (Collinson, 1996). Characters: fertile-ster-
ile leaf blade differentiation; absence of well-defined
sori; sporangia each with a transverse, subapical, contin-
uous annulus.
10. Family Lygodiaceae. — Climbing ferns.
single genus (Lygodium), ca. 25 spp.; monophyletic
(Skog & al., 2002; Wikström & al., 2002). Terrestrial,
pantropical. Characters: rhizomes creeping, slender, pro-
tostelic, bearing hairs; leaves indeterminate, climbing,
alternately pinnate; primary blade divisions (pinnae)
pseudodichotomously forking with a dormant bud in the
axils; veins free or anastomosing; sori on lobes of the
ultimate segments; sporangia abaxial, solitary, one per
sorus, each sporangium covered by an antrorse indusi-
um-like subtending flange; spores 128–256 per spo-
rangium, tetrahedral and trilete; gametophytes green,
cordate, surficial; x = 29, 30.
11. Family Anemiaceae. — Including Mohria-
One genus (Anemia, incl. Mohria), ca. 100+ spp.;
monophyletic (Skog & al., 2002; Wikström & al., 2002).
Terrestrial; primarily New World, but a few spp. in Afri-
ca, India, and islands in Indian Ocean. Characters: rhi-
zomes creeping to suberect, bearing hairs; leaves deter-
minate, mostly hemidimorphic or dimorphic; veins free,
dichotomous, occasionally casually anastomosing; spo-
rangia usually on a basal pair (sometimes more than two
pinnae, or all pinnae modified and fertile) of skele-
tonized, highly modified, often erect pinnae; spores
128–256 per sporangium, tetrahedral, with strongly par-
allel ridges (Tryon & Lugardon, 1991); gametophytes
green, cordate, surficial; x = 38.
12. Family Schizaeaceae.
Two genera (Ac-
tinostachys, Schizaea), ca. 30 spp.; monophyletic (Skog
& al., 2002; Wikström & al., 2002). The Cretaceous
Schizaeopsis is the oldest fossil assigned to this lineage
(Wikström & al., 2002). Terrestrial, pantropical. Charac-
ters: blades simple (linear) or fan-shaped, variously cleft
and with dichotomous free veins; sporangia on marginal,
elaminate, branched or unbranched projections at blade
tips, not in discrete sori, exindusiate; spores bilateral,
monolete, 128–256 per sporangium; gametophytes green
and filamentous (Schizaea), or subterranean and non-
green, tuberous (Actinostachys), and non-green; a puz-
zling array of base chromosome numbers: x = 77, 94,
I. ORDER SALVINIALES. — Water ferns, he-
terosporous ferns; incl. “Hydropteridales”, Marsil-
eales, Pilulariales.
Monophyletic (Hasebe & al., 1995;
Pryer, 1999; Pryer & al., 2001a, 2004b). Characters: fer-
tile-sterile leaf blade differentiation; veins anastomosing;
aerenchyma tissue often present in roots, shoots, and
petioles; annulus absent; plants heterosporous, spores
with endosporous germination; monomegaspory; game-
tophytes reduced.
13. Family Marsileaceae. — Clover ferns,
incl. Pilulariaceae.
Three genera (Marsilea, Pilularia,
Regnellidium), ca. 75 total spp.; monophyletic (Hasebe
& al., 1995; Pryer, 1999; Nagalingum & al., unpubl.).
Hennipman (1996) included both Salviniaceae and Azol-
laceae within Marsileaceae, but the spores of Marsileace-
ae differ markedly from those of Salviniaceae and Azol-
laceae (Schneider & Pryer, 2002). Rooted aquatics, in
ponds, shallow water, or vernal pools, with floating or
emergent leaf blades; subcosmopolitan. Characters:
stems usually long-creeping, slender, often bearing hairs;
leaflets 4, 2 or 0 per leaf; veins dichotomously branched
but often fusing toward their tips; sori borne in stalked
bean-shaped sporocarps (Nagalingum & al., in press),
these arising from the rhizomes or from the base of the
petioles, one to many per plant; heterosporous, micro-
spores globose, trilete, megaspores globose, each with an
acrolamella positioned over the exine aperture
(Schneider & Pryer, 2002); perine gelatinous; x=10
(Pilularia), 20 (Marsilea).
14. Family Salviniaceae. — Floating ferns,
mosquito ferns; incl. Azollaceae.
Two genera (Sal-
vinia, Azolla), ca. 16 spp.; monophyletic (Pryer & al.,
1995, 2004b; Reid & al., 2006). Some authors separate
the genera into two families (Schneller in Kubitzki,
Smith & al. • Fern classification55 (3) • August 2006: 705–731
1990). Plants free-floating, subcosmopolitan; fossil re-
cord beginning in Cretaceous (Collinson, 1996). Charac-
ters: roots present (Azolla) or lacking (Salvinia); stems
protostelic, dichotomously branched; leaves sessile, al-
ternate, small (ca. 1–25 mm long), round to oblong, en-
tire; veins free (Azolla) or anastomosing (Salvinia);
spores of two kinds (plants heterosporous), large mega-
spores and small microspores, these globose, trilete;
spore germination endosporic; x = 9 (Salvinia), the low-
est base chromosome number known in ferns, 22
Tree ferns; incl.
Dicksoniales, Hymenophyllopsidales, Loxomatales, Me-
taxyales, Plagiogyriales (Hasebe & al., 1995; Wolf & al.,
1999; Pryer & al., 2004b). Existing molecular evidence
indicates a close relationship among the included fami-
lies. The order is without obvious defining morphologi-
cal characters: some of the species have trunk-like stems
but others have creeping rhizomes; some have only hairs
on the stems and blades, others have scales; sori are ab-
axial or marginal, either indusiate or exindusiate; spores
are globose or tetrahedral-globose, with a trilete scar; ga-
metophytes green, cordate.
15. Family Thyrsopteridaceae. —
One genus,
Thyrsopteris, with a single species, T. elegans, endemic
to the Juan Fernández Islands; clearly related to tree
ferns, but of uncertain phylogenetic position within this
group (Korall & al., 2006). Characters: rhizomes as-
cending to erect, solenostelic, bearing runners, clothed
with stiff, pluricellular hairs; leaves large, 2–3.5 m long;
blades 3–5-pinnate, partially dimorphic (sori often
restricted to proximal segments); blade axes adaxially
grooved; veins free; sori terminal on the veins, the outer
and inner indusia fused to form asymmetric cup-like
structures, each sorus with a columnar, clavate recepta-
cle; sporangia with oblique annuli; spores globose-tetra-
hedral, with prominent angles; x = ca. 78.
16. Family Loxomataceae. —
Two genera (Lo-
xoma, Loxsomopsis), each with a single sp.; monophylet-
ic (Pryer & al., 2001a, 2004b; Lehnert & al., 2001;
Korall & al., 2006). South American Andes, southern
Central America, and New Zealand. Characters: rhi-
zomes long-creeping, solenostelic, bearing hairs with a
circular, multicellular base; blades bipinnate or more di-
vided; veins free, forked; indument of uniseriate (Loxso-
mopsis) to pluriseriate (Loxoma) bristles; sori marginal,
terminal on veins, each with an urceolate indusium and
elongate, often exserted receptacle; sporangia on thick,
short stalks, with a slightly oblique annulus; spores tetra-
hedral, trilete; gametophytes with scalelike hairs (occur-
ring also in some Cyatheaceae); x = 46 (Loxsomopsis),
50 (Loxoma).
17. Family Culcitaceae. —
One genus, Culcita,
with two species; monophyletic (Korall & al., 2006). Sis-
ter to Plagiogyriaceae, and not closely related to
Calochlaena, with which Culcita has often been associ-
ated. This separation is supported by anatomical charac-
ters (White & Turner, 1988; Schneider, 1996a). Terrestri-
al; Azores, Madeira, Tenerife, southwestern Europe, and
the Neotropics. Characters: rhizomes creeping or ascend-
ing, solenostelic, bearing articulate hairs; petioles in
cross-section each with gutter-shaped vascular bundle;
blades large, 4–5-pinnate-pinnatifid, sparingly hairy;
veins free, often forked; sori to 3 mm wide, terminal on
veins, paraphysate; outer indusia scarcely differentiated
from the laminar tissue, inner noticeably modified;
spores tetrahedral-globose, trilete; x = 66.
18. Family Plagiogyriaceae. —
A single genus
(Plagiogyria), with ca. 15 spp. (Zhang & Nooteboom,
1998); monophyletic (Korall & al., 2006). Characters:
stems creeping to usually erect, lacking hairs or scales;
leaves dimorphic; blades pectinate to 1-pinnate; veins
simple to 1-forked, free, or in fertile blades somewhat
anastomosing at their ends; young leaves densely cov-
ered with pluricellular, glandular, mucilage-secreting
hairs; sori exindusiate; sporangia borne on distal parts of
veins, seemingly acrostichoid; sporangial stalks 6-
rowed; annuli slightly oblique, continuous; spores tetra-
hedral, trilete; gametophytes green, cordate; x = 66?
19. Family Cibotiaceae
Korall, stat. nov. Based
on a full and direct reference to the Latin description
associated with subfam. Cibotioideae Nayar, Taxon 19:
234. 1970. – Type: Cibotium Kaulf., Jahrb. Pharm. 21:
53. 1820.
One genus (Cibotium), ca. 11 species; monophyletic,
with some affinity to Dicksoniaceae, as circumscribed
here (Korall & al., 2006). Terrestrial, amphipacific (east-
ern Asia, Malesia, Hawaii, southern Mexico and Central
America). Characters: rhizomes massive, creeping to
ascending or erect (to 6 m), solenostelic or dictyostelic,
bearing soft yellowish hairs at apices and persistent peti-
olar bases; fronds monomorphic, mostly 2–4 m long;
petioles hairy at bases, with three corrugated vascular
bundles arranged in an omega-shape; blades large, bipin-
nate to bipinnate-pinnatifid or more divided; secondary
and tertiary blade axes adaxially ridged; veins free, sim-
ple or forked to pinnate; stomata with 3 subsidiary cells;
sori marginal at vein ends, indusia bivalvate, each with a
strongly differentiated, non-green outer indusium and a
similarly modified tongue-like inner indusium, paraphy-
ses filiform; spores globose-tetrahedral, with prominent
angles and a well-developed equatorial flange; antheridi-
al walls 5-celled; x = 68.
The spores of Cibotiaceae are unlike those of all
other families in Cyatheales (Gastony, 1982; Tryon &
Lugardon, 1991), with a prominent equatorial flange, and
with usually thick, bold, ± parallel, sometimes anasto-
mosing ridges on the distal face; these ridges are the
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
result of a coarsely ridged exospore, which is overlain by
a thin, sometimes granulate perispore. The spores of
Lophosoria also have a prominent equatorial flange but
lack distal ridges (Tryon & Tryon, 1982; Tryon & Lu-
gardon, 1991). As far as is known, the chromosome base
number of x = 68 for Cibotium is also unique in the tree
fern clade. The Hawaiian species have been extensively
studied by Palmer (1994).
20. Family Cyatheaceae. —
Cyatheoids, scaly
tree ferns; incl. Alsophilaceae, Hymenophyllopsidaceae.
Ca. five genera: Alsophila (incl. Nephelea), Cyathea (incl.
Cnemidaria, Hemitelia, Trichipteris), Gymnosphaera,
Hymenophyllopsis, Sphaeropteris (incl. Fourniera);
600+ spp.; monophyletic, together with Dicksoniaceae,
Metaxyaceae, and Cibotiaceae constituting the “core tree
ferns” (Korall & al., 2006). Several studies have ad-
dressed relationships within Cyatheaceae (Conant & al.,
1995, 1996; Hasebe & al., 1995; Stein & al., 1996; Lantz
& al., 1999; Conant & Stein, 2001), and circumscriptions
of genera have varied widely (e.g., Tryon, 1970; Holttum
& Edwards, 1983). Several of these studies show con-
vincingly that Cnemidaria nests within Cyathea (Conant
& al., 1995; Conant & Stein, 2001; Korall & al., 2006).
Hennipman (1996) included all other families here
placed in Cyatheales (excepting Hymenophyllopsidace-
ae, unplaced in his “consensus” classification) in his Cy-
atheaceae. Hymenophyllopsis (ca. eight species) has thin
leaves lacking stomates, and is confined to the sandstone
tepuis of the Guayana Shield (eastern Venezuela, Guya-
na, northern Brazil). It has nearly always been regarded
as an isolated genus in its own family (e.g., by Copeland,
1947; Tryon & Tryon, 1982). In the analysis by Wolf &
al. (1999), a close relationship of Hymenophyllopsis to
Cyatheaceae was suggested, based on a small taxonomic
sampling. A larger sampling by Korall (in prep.) indi-
cates that Hymenophyllopsis, as well as Cnemidaria and
Trichipteris, all nest within Cyathea, and together form a
well supported neotropical clade. The spores of Hymeno-
phyllopsis are remarkably similar to some species of
Cyathea (compare, e.g., Figs. 14.8–14.11 with 26.15–
26.18 in Tryon & Tryon, 1982). Characters associating
Hymenophyllopsis with Cyatheaceae include the pres-
ence of true scales on the rhizomes, petiole bases, and
sometimes on the blades. Mostly arborescent, pantropi-
cal; fossils beginning in Jurassic or early Cretaceous.
Characters: stems with polycyclic dictyosteles, apices
(and usually petiole bases) covered with large scales,
sometimes also with trichomidia (scurf = small scales) or
hairs; leaves usually large (to 5 m); petioles with obvi-
ous, usually discontinuous pneumathodes in two lines;
blades 1–3-pinnate (rarely simple); veins simple to
forked, free, rarely anastomosing (mostly in some Cy-
athea); sori superficial (abaxial) or terminal on the veins
and marginal or submarginal (Hymenophyllopsis), round,
exindusiate, or indusia saucer-like, cup-like, or globose
and completely surrounding sporangia, or bivalvate
(Hymenophyllopsis); sporangia maturing gradately, with
oblique annuli; receptacles raised; paraphyses usually
present; spores tetrahedral, trilete, variously ornamented;
gametophytes green, cordate; x = 69 (Hymenophyllopsis
not yet counted).
21. Family Dicksoniaceae,
nom. cons.
(Dicksonioids; incl. Lophosoriaceae). Three genera: Cal-
ochlaena, Dicksonia, Lophosoria). Ca. 30 spp.; mono-
phyletic (Korall & al., 2006). Terrestrial; eastern Asia,
Australasia, Neotropics, St. Helena. Characters: mostly
arborescent or with erect or ascending rhizomes; rhi-
zomes with polycyclic dictyosteles, or solenostelic
(Calochlaena); stem apices and usually petiolar bases
covered with uniseriate hairs; blades large, 2–3 pinnate;
veins simple to forked, free; sori abaxial and exindusiate
(Lophosoria) or marginal (Calochlaena, Dicksonia) and
each with a bivalvate or cup-like indusium, the adaxial
(outer) valve formed by the reflexed segment margin and
often differently colored; sporangia with oblique annuli;
receptacles raised; paraphyses often present, filiform;
spores globose or tetrahedral, trilete; x = 56
(Calochlaena), 65 (Dicksonia, Lophosoria).
Lophosoria (3 spp.) is distinctive in having spores
with a prominent subequatorial flange, with the proximal
face coarsely tuberculate, the distal face perforate. It has
often been recognized as comprising its own family,
Lophosoriaceae (Pichi Sermolli, 1977; Tryon & Tryon,
1982; Kramer in Kubitzki, 1990).
22. Family Metaxyaceae. —
A single genus (Me-
taxya), 2 spp.; monophyletic (Smith & al., 2001). Terres-
trial, Neotropics. Characters: rhizomes short-creeping to
ascending, dorsiventral, solenostelic, apices covered
with pluricellular hairs; petioles each with an omega-
shaped, corrugated vascular bundle; blades simply pin-
nate; veins free, simple or forked at the base, ± parallel;
sori abaxial, round, scattered in several ill-defined rows,
often with several sori on the same vein, with numerous
filiform paraphyses, exindusiate; sporangia maturing
simultaneously; sporangial stalks 4-rowed; annuli verti-
cal or slightly oblique; spores 64 per sporangium, glo-
bose, trilete; x = 95, 96.
Including “As-
pidiales”, Aspleniales, Athyriales, Blechnales, “Davallia-
les”, Dennstaedtiales, Dryopteridales, Lindsaeales, Lon-
chitidales, Monachosorales, Negripteridales, Parkeriales,
Platyzomatales, Pteridales, Saccolomatales, Thelypteri-
dales. Monophyletic (Hasebe & al., 1995; Pryer & al.,
1995, 2001a, 2004b; Schneider & al., 2004c). Charac-
ters: indusia laterally or centrally attached (indusia lost in
many lineages); sporangial stalks 1–3 cells thick, often
long; sporangial maturation mixed; sporangia each with
a vertical annulus interrupted by the stalk and stomium;
Smith & al. • Fern classification55 (3) • August 2006: 705–731
gametophytes green, usually cordate (sometimes ribbon-
shaped in some epiphytes), surficial.
23. Family Lindsaeaceae. —
Lindsaeoids; incl.
Cystodiaceae, Lonchitidaceae. Ca. eight genera: Cystodi-
nomeris, Tapeinidium, Xyropteris
; in an unpublished
thesis, Barcelona (2000) advocated the establishment of
three additional genera allied to Odontosoria and Sphe-
nomeris. Ca. 200 spp.; most likely monophyletic (Wolf
& al., 1994; Pryer & al., 2004b; Korall & al., in press;
Schuettpelz & al., in press). The inclusion of Lonchitis
(traditionally associated with dennstaedtioid ferns) in
Lindsaeaceae is puzzling on morphological grounds, but
molecular evidence strongly suggests it belongs with the
lindsaeoid ferns. Terrestrial, or infrequently epipetric or
epiphytic, pantropical. Characters: roots with sclerenchy-
matous outer cortex combined with an innermost cortical
layer six cells wide (Schneider, 1996a) (excepting Lon-
chitis and Cystodium); rhizomes short- to long-creeping,
protostelic with internal phloem, or in a few taxa soleno-
stelic, bearing generally narrow, basally attached, non-
clathrate scales or uniseriate hairs; blades 1–3-pinnate or
more divided, generally glabrous; veins usually free, for-
king, occasionally anastomosing, without included vein-
lets; sori marginal or submarginal, indusiate, indusia open-
ing towards the margin (extrorse), sometimes also at-
tached at the sides, or sori covered by the reflexed seg-
ment margin (Lonchitis); spores tetrahedral, trilete, infre-
quently bilateral, monolete; gametophytes green, cor-
date; x = 34, 38, 39, 44, 47, 48, 49, 50, 51, perhaps oth-
The position of Cystodium is clearly among Poly-
podiales, and not Dicksoniaceae (in Cyatheales), where it
has historically been placed, e.g., by Kramer in Kubitzki,
1990; Stevenson & Loconte, 1996 (Korall & al., in
press). Croft (1986) discussed its differences from dick-
sonioids and recognized it at family rank. A relationship
to other lindsaeoids is suggested by the molecular evi-
dence, and this is reflected in our classification.
However, expanded taxon sampling within early-diverg-
ing lineages of Polypodiales is necessary to confirm this
or to determine whether recognition of a monotypic fam-
ily Cystodiaceae is warranted (Korall & al., in press).
24. Family Saccolomataceae. —
One genus, ca.
12 spp.; apparently monophyletic, but more sampling is
needed to determine whether the Old World species are
congeneric with those from the New World. The rela-
tionships of Saccoloma (incl. Orthiopteris) have been
contentious. Kramer (in Kubitzki, 1990) treated Saccolo-
ma as a subfamily within Dennstaedtiaceae. Terrestrial,
pantropical. Characters: rhizomes short-creeping to erect
and somewhat trunk-like (long-creeping in most Lind-
saeaceae and Dennstaedtiaceae) and dictyostelic (usually
solenostelic in Dennstaedtiaceae, protostelic with inter-
nal phloem in Lindsaeaceae); petioles each with an
omega-shaped vascular strand (open end adaxial); blades
pinnate to decompound, lacking articulate hairs (as found
in Dennstaedtiaceae); veins free; sori terminal on the
veins, indusia pouch- or cup-shaped; spores globose-
tetrahedral, surface with distinctive ± parallel, branched
ridges; x = ca. 63.
25. Family Dennstaedtiaceae. —
ioids; incl. Hypolepidaceae, Monachosoraceae, Pteridia-
ceae. Ca. 11 genera: Blotiella, Coptodipteris, Denn-
staedtia (incl. Costaricia
), Histiopteris, Hypolepis,
Leptolepia, Microlepia, Monachosorum, Oenotrichia
, Paesia, Pteridium (bracken). Ca. 170 spp.; mono-
phyletic, if lindsaeoid ferns are excluded (Pryer & al.,
2004b; Schuettpelz & al., in press). Monachosoraceae
nests within Dennstaedtiaceae (Wolf & al., 1994; Wolf,
1995, 1997; Pryer & al., 2004b; Schuettpelz & al., 2006).
Terrestrial, sometimes scandent; pantropical. Characters:
rhizomes mostly long-creeping, often siphonostelic or
polystelic, bearing jointed hairs; petioles often with
epipetiolar buds, usually with a gutter-shaped vascular
strand (open end adaxial); blades often large, 2–3-pin-
nate or more divided; indument of hairs; veins free,
forked or pinnate, rarely anastomosing and then without
included veinlets; sori marginal or submarginal, linear or
discrete, indusia linear or cup-like at blade margins, or
reflexed over sori; spores tetrahedral and trilete, or reni-
form and monolete; gametophytes green, cordate; x = 26,
29, 30, 31, 33, 34, 38, 46, 47, 48, and probably others.
26. Family Pteridaceae. —
Pteroids or pterido-
ids; incl. Acrostichaceae, Actiniopteridaceae, Adiantace-
ae (adiantoids, maidenhairs), Anopteraceae, Antrophya-
ceae, Ceratopteridaceae, Cheilanthaceae (cheilanthoids),
Cryptogrammaceae, Hemionitidaceae, Negripteridaceae,
Parkeriaceae, Platyzomataceae, Sinopteridaceae, Taeniti-
daceae (taenitidoids), Vittariaceae (vittarioids, shoestring
ferns). Ca. 50 genera, 950 spp. Constituent genera, some
of them notoriously polyphyletic or paraphyletic and in
need of redefinition (e.g., Cheilanthes), include Acrosti-
chum, Actiniopteris, Adiantopsis, Adiantum, Aleur-
itopteris, Ananthacorus, Anetium, Anogramma, Antro-
phyum, Argyrochosma, Aspidotis, Astrolepis, Austro-
gramme, Bommeria, Cassebeera, Ceratopteris, Cero-
, Cheilanthes, Cheiloplecton, Coniogramme,
Cosentinia (Nakazato & Gastony, 2001), Cryptogram-
ma, Doryopteris, Eriosorus, Haplopteris, Hecistopteris,
Hemionitis, Holcochlaena, Jamesonia, Llavea, Mildella,
Monogramma, Nephopteris
, Neurocallis, Notholaena,
Ochropteris, Onychium, Paraceterach, Parahemionitis,
Pellaea (Kirkpatrick, unpubl.), Pentagramma,
Pityrogramma, Platyloma, Platyzoma, Polytaenium,
Pteris (incl. Afropteris, Anopteris), Pterozonium,
Radiovittaria, Rheopteris, Scoliosorus, Syngramma,
Taenitis, Trachypteris, and Vittaria. The family thus
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
defined is monophyletic (Gastony & Rollo, 1995, 1998;
Hasebe & al., 1995; Pryer & al., 1995; Gastony &
Johnson, 2001; Schneider & al., 2004c; Zhang & al.,
2005). Pteridaceae comprises five monophyletic groups,
and if it were to be formally subdivided to reflect this, at
either family or subfamily rank, the following taxa could
be recognized: (1) Parkeriaceae, or Parkerioideae
(Acrostichum and Ceratopteris), (2) Adiantaceae, or
Adiantoideae (Adiantum and the ten vittarioid genera;
Crane & al., 1995; Hasebe & al., 1995; Hennipman,
1996; Crane, 1997; Huiet & Smith, unpubl.); (3)
Cryptogrammaceae (comprising Coniogramme, Crypto-
gramma, and Llavea; Zhang & al., 2005), no subfamily
name available; (4) Sinopteridaceae, or Cheilan-
thoideae); and (5) Pteridaceae s.s., or Pteridoideae, con-
taining Pteris (perhaps not monophyletic), its immediate
allies, and the taenitioid ferns (Taenitis and allies;
Sánchez-Baracaldo, 2004a, b).
Terrestrial, epipetric, or epiphytic, subcosmopolitan,
but most numerous in tropics and arid regions.
Characters: rhizomes long- to short-creeping, ascending,
suberect, or erect, bearing scales (less often, only hairs);
blades monomorphic, hemidimorphic, or dimorphic in a
few genera, simple (mostly vittarioids), pinnate, or some-
times pedate, sometimes decompound; veins free and
forking, or variously anastomosing and forming a reticu-
late pattern without included veinlets; sori marginal or
intramarginal, lacking a true indusium, often protected
by the reflexed segment margin, or sporangia along the
veins; sporangia each with a vertical, interrupted annu-
lus, receptacles not or only obscurely raised; spores glo-
bose or tetrahedral, trilete, variously ornamented; mostly
x = 29, 30.
Platyzoma, sometimes recognized as an isolated
family, is aberrant in chromosome base number (x = 38;
Tindale & Roy, 2002) and in having dimorphic spores
(so-called “incipient heterospory”; Tryon, 1964), but
nests with other genera of Pteridaceae, subfam.
Pteridoideae (Hasebe & al., 1995; Pryer & al., 1995).
Ceratopteris (3 spp., monophyletic) nests within
Pteridaceae in all molecular analyses, and it appears to be
sister to Acrostichum (Hasebe & al., 1995; Pryer & al.,
1995). It has a number of strong autapomorphies that
separate it from other Pteridaceae: coarsely ridged spores
with parallel striations; spores 32 or fewer per spo-
rangium; sporangia with ill-defined annuli; aquatic habi-
tat; x = 38. Consequently, many taxonomists have placed
it in its own family, Parkeriaceae (e.g., Copeland, 1947;
Pichi Sermolli, 1977). Many of these autapomorphies
(reduced spore number, loss of annulus) are probably a
consequence of the shift to aquatic habitats.
The vittarioid genera include: Ananthacorus, Aneti-
um, Antrophyum, Haplopteris, Hecistopteris, Mono-
gramma, Polytaenium, Radiovittaria, Rheopteris, Scoli-
osorus, and Vittaria. The presence of Rheopteris in this
clade is now clear (Huiet & Smith, unpubl.). Characters
include linear, mostly simple blades, sori along veins or
in linear grooves, clathrate stem scales; presence of “spi-
cular” cells in blade epidermis (shared with a few genera
of Pteridaceae, e.g., Adiantum). Spores in the vittarioid
ferns are predominantly trilete, but reversals to the
monolete condition have occurred in Vittaria.
27. Family Aspleniaceae. —
spleenworts. From one to ten genera (generic delimita-
tion in doubt, in light of all recent molecular data, e.g.,
van den Heede & al., 2003; Schneider & al., 2004b,
2005; Perrie & Brownsey, 2005). Regardless of the clas-
sification adopted, a huge majority of the species are in
Asplenium, even if that genus is construed in a fairly
strict sense; the segregate genera Camptosorus and Loxo-
scaphe clearly nest within Asplenium s.l., or appear relat-
ed to species heretofore generally placed in Asplenium
(Murakami & al., 1999; Gastony & Johnson, 2001;
Schneider & al., 2004b). So also are the generic segre-
gates Diellia (endemic to Hawaii), Pleurosorus, Phylli-
tis, Ceterach, Thamnopteris, and several others little
used in recent years (Murakami & al., 1999; Pinter & al.,
2002; van den Heede & al., 2003; Schneider & al.,
2004b). One expects that the oft-recognized, but still un-
sampled genera Antigramma
, Holodictyum
, Schaffner-
, and Sinephropteris
also nest in Asplenium. Hymen-
asplenium, however, with a different chromosome base
number than nearly all of the other segregates, as well as
distinct root characters (Schneider, 1996b; Schneider &
al., 2004b), appears to represent the sister clade to the
rest of the species in the family, and this name could be
adopted as a well-supported segregate genus. Ca. 700+
spp; monophyletic (Murakami & Schaal, 1994; Hasebe
& al., 1995; Murakami & al., 1999; Gastony & Johnson,
2001; van den Heede & al., 2003; Schneider & al.,
2004b, 2005; Perrie & Brownsey, 2005).
Terrestrial, epipetric, or epiphytic, subcosmopolitan,
but most numerous in the tropics. Characters: rhizomes
creeping, ascending, or suberect, bearing clathrate scales
at apices and petiole bases (and sometimes other axes);
petioles with back-to-back C-shaped vascular strands,
these fusing distally into an X-shape; blades monomor-
phic, usually lacking acicular hairs on axes and/or lami-
na, often with microscopic clavate hairs; veins pinnate or
forking, usually free, infrequently reticulate and then
without included veinlets; sori elongate (linear) along the
veins, not usually back-to-back on the same vein, usual-
ly with laterally attached, linear indusia; sporangial
stalks long, 1-rowed; spores reniform, monolete, with a
decidedly winged perine; x = 36 (mostly), but x = 38, 39
in Hymenasplenium (Murakami, 1995), 38 in Boniniella.
28. Family Thelypteridaceae. —
or thelypteridoids; incl. “Sphaerostephanaceae”. Circa
Smith & al. • Fern classification55 (3) • August 2006: 705–731
5–30 genera, depending on taxonomic viewpoint: com-
monly accepted segregates are Cyclosorus (incl. Ampel-
, Amphineuron
, Chingia,Christella, Cyclogram-
, Cyclosorus s.s., Glaphyropteridopsis, Goniopteris,
Meniscium, Menisorus
, Mesophlebion, Pelazoneuron,
Plesioneuron, Pneumatopteris, Pronephrium, Pseudo-
cyclosorus, Sphaerostephanos, Stegnogramma, Steirop-
teris, Trigonospora), Macrothelypteris, Phegopteris,
Pseudophegopteris, and Thelypteris (incl. Amauropelta,
Coryphopteris, Metathelypteris, Oreopteris, Parathelyp-
teris, and Thelypteris s.s.) (see Holttum, 1971; Smith &
Cranfill, 2002). Ca. 950 spp; monophyletic (Hasebe &
al., 1995; Smith & Cranfill, 2002; Yatabe & al., 2002).
Hennipman (1996) also included Blechnaceae and the
athyrioid ferns in this family, a definition that would
make Thelypteridaceae difficult or impossible to define
Terrestrial, rarely epipetric, pantropical, a few tem-
perate. Characters: rhizomes creeping, ascending, or
erect, bearing scales at apices, these non-clathrate, usual-
ly bearing acicular hairs; petioles in cross-section with
two elongate or crescent-shaped vascular bundles facing
one another, these uniting distally into a gutter-shape;
blades monomorphic or occasionally dimorphic, usually
pinnate or pinnate-pinnatifid; veins pinnate, free to vari-
ously and usually very regularly anastomosing, with or
without included veinlets; indument of acicular hyaline
hairs on blades and rhizome scales; sori abaxial, round to
oblong, rarely elongate along veins, with reniform indu-
sia or exindusiate; sporangia with 3-rowed, short to long
stalks; spores ellipsoid, monolete, perine winged to spin-
ulose; x = 27–36. Indusia have been lost independently in
many lineages within the family.
29. Family Woodsiaceae. —
Athyrioids, lady
ferns; incl. Athyriaceae, Cystopteridaceae. Circa 15 gen-
era as defined here, ca. 700 spp., nearly 85% of them in
the two main genera, Athyrium and Diplazium (incl.
Callipteris, Monomelangium), which are both probably
paraphyletic (Wang & al., 2003). Other widely recog-
nized genera include Acystopteris, Cheilanthopsis, Cor-
nopteris, Cystopteris, Deparia (incl. Lunathyrium, Dryo-
athyrium, Athyriopsis, and Dictyodroma; Sano & al.,
2000b), Diplaziopsis, Gymnocarpium (incl. Currania),
Hemidictyum, Homalosorus, Protowoodsia
, Pseudocys-
topteris, Rhachidosorus, and Woodsia (incl. Hymenocys-
; see Shmakov, 2003). This family has been variously
circumscribed, and its limits are still uncertain (Hasebe
& al., 1995; Sano & al., 2000a, b). Wang & al. (2004) di-
vided the Athyriaceae (excluding woodsioid ferns), by
far the largest component in the family, into five subfam-
ilies: Cystopteroideae, Athyrioideae, Deparioideae, Di-
plazioideae, and Rhachidosoroideae. As delimited here,
the Woodsiaceae may be paraphyletic to the Aspleniace-
ae, Blechnaceae + Onocleaceae, and Thelypteridaceae,
but support for this paraphyly—or alternatively for the
monophyly of the family as here defined—is lacking in
broad analyses (Hasebe & al., 1995; Sano & al., 2000a;
Schneider & al., 2004c). Because of this uncertainty,
combined with the morphological grounds for the recog-
nition of the Woodsiaceae as here circumscribed, we
believe it is premature to adopt the alternative of erecting
(or resurrecting) numerous small families to house its
constituent genera. Further sampling will likely shed
additional light on this subject, and the recognition of
several additional families may be warranted.
Mostly terrestrial, subcosmopolitan. Characters: rhi-
zomes creeping, ascending, or erect; scales at apices,
these usually non-clathrate, glabrous, glandular, or cili-
ate; petioles with two elongate or crescent-shaped vascu-
lar bundles facing one another, these uniting distally into
a gutter-shape; blades monomorphic, rarely dimorphic;
veins pinnate or forking, free, uncommonly anastomos-
ing and then without included veinlets; sori abaxial,
round, J-shaped, or linear with reniform to linear indusia,
or exindusiate; spores reniform, monolete, perine
winged, ridged or spiny; mostly x = 40, 41, also 31
(Hemidictyum), 33, 38, 39 (Woodsia), and 42 (Cysto-
30. Family Blechnaceae. —
Blechnoids; incl.
Stenochlaenaceae). Currently ca. nine genera recognized
(Blechnum s.l., Brainea, Doodia, Pteridoblechnum, Sad-
leria, Salpichlaena, Steenisioblechnum, Stenochlaena,
Woodwardia). Most of the existing recognized genera
nest within Blechnum s.l., and their acceptance is depen-
dent upon a revised recircumscription of Blechnum s.l.,
which is manifestly paraphyletic in its current usage
(Nakahira, 2000; Cranfill, 2001). Ca. 200 spp.; mono-
phyletic, sister to Onocleaceae (Hasebe & al., 1995;
Cranfill, 2001; Cranfill & Kato, 2003). Woodwardia
(incl. Anchistea, Chieniopteris, Lorinseria) appears to be
an early-branching member of the Blechnaceae (Cranfill
& Kato, 2003). Characters: rhizomes creeping, ascend-
ing, or erect, sometimes trunk-like, often bearing stolons,
scaly at apex (and on blades), scales non-clathrate; peti-
oles with numerous, round, vascular bundles arranged in
a ring; leaves monomorphic or often dimorphic; veins
pinnate or forking, free to variously anastomosing, are-
oles without included veinlets, on fertile leaves forming
costular areoles bearing the sori; sori in chains or linear,
often parallel and adjacent to midribs, indusiate, with lin-
ear indusia opening inwardly (toward midribs); sporan-
gia with 3-rowed, short to long stalks; spores reniform,
monolete, perine winged; gametophytes green, cordate; x
= 27, 28, 31–37 (Blechnum and segregates, Woodward-
ia); 40 (Salpichlaena).
31. Family Onocleaceae. —
Onocleoids. Four
genera: Matteuccia, Onoclea, Onocleopsis, Pentarhizidi-
um. 5 spp.; monophyletic, sister to Blechnaceae (Hasebe
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
& al., 1995; Gastony & Ungerer, 1997). Family circum-
scription follows Pichi Sermolli (1977) and Gastony and
Ungerer (1997, their tribe Onocleeae of Dryopteridace-
ae). Terrestrial, largely in north-temperate regions; Char-
acters: rhizomes long- to short-creeping to ascending,
sometimes stoloniferous (Matteuccia and Onocleopsis);
leaves strongly dimorphic; petioles with two vascular
bundles uniting distally into a gutter-shape; blades pin-
natifid or pinnate-pinnatifid; veins free or anastomosing,
lacking included veinlets; spores reniform, brownish to
green; sori enclosed (sometimes tightly) by reflexed lam-
inar margins, also with membranous, often fugacious
true indusia; x = 37 (Onoclea), 39, 40 (other genera).
32. Family Dryopteridaceae. —
or dryopteridoids; incl. “Aspidiaceae”, Bolbitidaceae,
Elaphoglossaceae, Hypodematiaceae, Peranemataceae.
Circa 40–45 genera, 1700 spp., of which 70% are in four
genera (Ctenitis, Dryopteris, Elaphoglossum, and Poly-
stichum). Genera include Acrophorus, Acrorumohra,
Adenoderris, Arachniodes, Ataxipteris
, Bolbitis (incl.
Egenolfia), Coveniella
, Ctenitis, Cyclodium, Cyrtogo-
nellum, Cyrtomidictyum, Cyrtomium (Lu & al., 2005),
Didymochlaena, Dryopolystichum
, Dryopsis, Dryo-
pteris (incl. Nothoperanema; Geiger & Ranker, 2005),
Elaphoglossum (incl. Microstaphyla
, Peltapteris;
Rouhan & al., 2004; Skog & al., 2004), Hypodematium,
Lastreopsis, Leucostegia, Lithostegia
, Lomagramma,
Maxonia, Megalastrum, Oenotrichia p.p. (Tindale &
Roy, 2002), Olfersia, Peranema, Phanerophlebia,
Polystichum (incl. Papuapteris, Plecosorus; Little &
Barrington, 2003), Polybotrya, Polystichopsis, Revwat-
(Tindale & Roy, 2002), Rumohra, Stenolepia
Stigmatopteris, and Teratophyllum.
Terrestrial, epipetric, hemiepiphytic, or epiphytic,
pantropical, also with many temperate representatives.
Characters: rhizomes creeping, ascending or erect, some-
times scandent or climbing, with non-clathrate scales at
apices; petioles with numerous round, vascular bundles
arranged in a ring; blades monomorphic, less often
dimorphic, sometimes scaly or glandular, uncommonly
hairy; veins pinnate or forking, free to variously anas-
tomosing, with or without included veinlets; sori usually
round, indusia round-reniform or peltate (lost in several
lineages), or sori exindusiate, acrostichoid in a few line-
ages; sporangia with 3-rowed, short to long stalks; spores
reniform, monolete, perine winged; x = 41 (nearly all
genera counted), rarely 40 (presumably derived).
Dryopteridaceae, as defined here, is almost certainly
monophyletic, if Didymochlaena, Hypodematium, and
Leucostegia are excluded (Hasebe & al., 1995; Tsutsumi
& Kato, 2006). The inclusion of these three genera may
render this family paraphyletic, but they are tentatively
included here pending further studies to address their
precise phylogenetic affinities. Didymochlaena, with a
single species, has generally been associated with other
members of the Dryopteridaceae (as here defined). The
three closely related species of Hypodematium, on the
other hand, have been variously treated: as composing a
monogeneric family Hypodematiaceae; as allied to the
athyrioid ferns (e.g., by Kramer in Kubitzki, 1990, pre-
sumably based on the presence of two vascular bundles
in the petiole bases); or as close to Dryopteris (e.g.,
Tryon & Lugardon, 1991, using evidence from spore
morphology). Leucostegia is nearly always placed in
Davalliaceae (e.g., by Kramer, 1990), because of its sim-
ilar indusia and sori terminal on the veins, but it differs
from members of Davalliaceae in the terrestrial habit, the
more strongly verrucate spores with rugulate perispore
(Tryon & Lugardon, 1991), and x = 41 (vs. x = 40 in Da-
valliaceae). In a molecular phylogenetic analysis by
Schneider & al. (2004c), Didymochlaena and Hypode-
matium were resolved as sister to one another, and
together sister to the remainder of the Eupolypods I clade
(Fig. 2), but support for these relationships was lacking.
Tsutsumi & Kato (2006) found support for a sister rela-
tionship between Hypodematium and Leucostegia, and
also support for these as sister to the remaining Eu-
polypods I, but Didymochlaena was unsampled in their
analysis. Based on these results, we therefore believe it
would be premature to segregate these genera from the
Within Dryopteridaceae, as defined here, nests
Elaphoglossum (Hasebe & al., 1995; Sano & al., 2000a).
Sometimes it is included in its own family Elaphoglos-
saceae, e.g., by Pichi Sermolli (1977), with 600–800
spp., many still undescribed. Elaphoglossaceae was re-
garded as comprising three genera by Pichi Sermolli
(1977), but Microstaphyla and Peltapteris nest within
Elaphoglossum (
phoglossum.html; Mickel & Atehortúa, 1980; Rouhan &
al., 2004; Skog & al., 2004). Relationships of Elapho-
glossum are often considered to be with Lomariopsis
(Kramer in Kubitzki, 1990), but this is refuted by two un-
published topologies. Elaphoglossaceae, narrowly de-
fined, is monophyletic (Skog & al., 2001, 2004), but to
exclude it from Dryopteridaceae s.s., as delimited above,
renders the latter paraphyletic. Characters of Elaphoglos-
sum include simple blades (usually) and dimorphic
leaves with acrostichoid sporangia.
Several authors have treated most of the genera with-
in our concept of Dryopteridaceae, as well as Tectariace-
ae, Woodsiaceae, and Onocleaceae, as comprising a
much larger family Dryopteridaceae s.l., with slightly
varying circumscriptions (e.g., Tryon & Tryon, 1982;
Kramer in Kubitzki, 1990; Wagner & Smith, 1993). With
such a broad circumscription, and unless several other
well-circumscribed families (e.g., Aspleniaceae, Blech-
naceae, Polypodiaceae, Thelypteridaceae) are included,
Smith & al. • Fern classification55 (3) • August 2006: 705–731
Dryopteridaceae is certainly paraphyletic.
33. Family Lomariopsidaceae. —
sids; incl. Nephrolepidaceae, sword ferns. Four genera:
Cyclopeltis, Lomariopsis, Nephrolepis, and Thysanosori-
; ca. 70 species. Characters: rhizomes creeping or
sometimes climbing (plants hemiepiphytic); petioles
with round vascular bundles arranged in a gutter-shape;
blades 1-pinnate, pinnae entire or crenate, often articu-
late, auriculate in some genera; veins free, ± parallel or
pinnate; sori discrete, round, and with round-reniform to
reniform indusia, or exindusiate, or sporangia acrosti-
choid and the fronds dimorphic; spores bilateral, mono-
lete, variously winged or ornamented; x = 41 (lower
numbers known in some Lomariopsis species).
Based on published and unpublished results, it ap-
pears likely that these genera form a monophyletic
group, despite the fact that such an assemblage has never
been proposed. Lomariopsidaceae (sensu Kramer in
Kubitzki, 1990; Moran in Davidse & al., 1995) was con-
strued to comprise six genera (containing ca. 800+
species): Bolbitis (and segregates Edanyoa, Egenolfia),
Elaphoglossum, Lomagramma, Lomariopsis, Terato-
phyllum, and Thysanosoria
. Based on available evi-
dence, we place all of the aforementioned genera except
Lomariopsis (and Thysanosoria, which lacks molecular
data, but appears to be closely related to Lomariopsis) in
the Dryopteridaceae (see above). Nephrolepis, with ca.
20 spp., has sometimes been included in a monogeneric
family Nephrolepidaceae (Kramer in Kubitzki, 1990). It
has been resolved and supported as sister to a large clade
comprising the Tectariaceae, Oleandraceae, Polypodia-
ceae, and Davalliaceae (Hasebe & al., 1995; Schneider &
al., 2004c); however, Lomariopsis was not included in
these analyses. When Lomariopsis is included, Nephrole-
pis is resolved as sister to it, and these two genera, in
turn, are strongly supported as sister to the aforemen-
tioned larger clade (Tsutsumi & Kato, 2006) and there-
fore to be expunged from the Dryopteridaceae. Although
we have here decided to tentatively include Nephrolepis
in the Lomariopsidaceae, the monophyly of this clade
requires additional scrutiny, and thus Nephrolepidaceae
may eventually require recognition.
34. Family Tectariaceae. —
Tectarioids; incl.
“Dictyoxiphiaceae”, “Hypoderriaceae”. 8–15 genera:
, Arthropteris, Heterogonium, Hypoder-
, Pleocnemia, Psammiosorus, Psomiocarpa
, Pteri-
drys, Tectaria s.l. (incl. Amphiblestra
, Cionidium, Ctenitopsis, Dictyo-
xiphium, Fadyenia, Hemigramma, Pleuroderris
, Pseu-
, Quercifilix, and perhaps other genera men-
tioned above), and Triplophyllum (Holttum, 1986); ca.
230 species, most in Tectaria s.l. Generic limits, espe-
cially within Tectaria s.l., are still very much in doubt.
The family appears monophyletic with the definition
given. Including Tectariaceae within an expanded
Dryopteridaceae renders the latter polyphyletic. Ctenitis,
Lastreopsis, and several other genera here included in
Dryopteridaceae have often been considered closely re-
lated to tectarioid ferns (Pichi Sermolli, 1977; Holttum,
1986; Moran in Davidse & al., 1995), but molecular data
suggest otherwise (Hasebe & al., 1995). Terrestrial,
pantropical. Characters: rhizomes usually short-creeping
to ascending, dictyostelic, bearing scales; petioles not
abscising, with a ring of vascular bundles in cross-sec-
tion; blades simple, pinnate, or bipinnate, sometimes de-
compound; indument of jointed, usually short stubby
hairs on the axes, veins, and sometimes laminar tissue,
especially on rachises and costae adaxially; veins free or
often highly anastomosing, sometimes with included
veinlets; indusia reniform or peltate (lost in several line-
ages); spores brownish, reniform, monolete, variously
ornamented; x = 40 (a few genera with x = 41, some dys-
ploids with x = 39).
Arthropteris is apparently not closely related to
Oleandra, as previously suggested (Kramer in Kubitzki,
1990), nor to Nephrolepis, as suggested by Pichi Sermol-
li (1977). Analyses that have included it show it to be sis-
ter to tectarioid ferns (Hasebe & al., 1995; Tsutsumi &
Kato, 2006). Psammiosorus, a monotypic genus endem-
ic to Madagascar, has in turn been placed close to Ar-
thropteris (Kramer, in Kubitzki, 1990) or even within Ar-
thropteris (Tryon & Lugardon, 1991, on the basis of the
spore ornamentation). Therefore, both Arthropteris and
Psammiosorus are tentatively assigned to Tectariaceae,
although a Tectariaceae that includes them is more diffi-
cult to define morphologically.
35. Family Oleandraceae. —
Monogeneric, ca.
40 spp., sister to Davalliaceae + Polypodiaceae (Hasebe
& al., 1995; Schneider & al., 2004; Tsutsumi & Kato,
2006). Kramer (in Kubitzki, 1990), included two genera
in addition to Oleandra: Arthropteris (ca. 12 spp.), and
Psammiosorus (monotypic), but with this broader cir-
cumscription, the family is clearly polyphyletic; we
include both of these genera in Tectariaceae. Species are
terrestrial, epilithic or often secondary hemiepiphytes.
Characters: blades simple; leaves articulate, abscising
cleanly upon senescence from pronounced phyllopodia;
sori indusiate, indusia round-reniform; spores reniform,
monolete; x = 41.
36. Family Davalliaceae. —
Davallioids; excl.
Gymnogrammitidaceae. 4–5 genera: Araiostegia, Daval-
lia (incl. Humata, Parasorus, Scyphularia), Davallodes,
Pachypleuria; ca. 65 spp. Monophyletic, sister to Poly-
podiaceae (Hasebe & al., 1995; Ranker & al., 2004;
Schneider & al., 2004d; Tsutsumi & Kato, 2005), but
more information needed. Gymnogrammitis and Leuco-
stegia are often included in Davalliaceae but the former
belongs in Polypodiaceae (Schneider & al., 2002b),
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
while the latter is seemingly allied to Hypodematium
(Dryopteridaceae; Tsutsumi & Kato, 2005). Generic lim-
its of Araiostegia, Davallia, and Pachypleuria relative to
each other are ill-defined, and all of these genera appear
to be paraphyletic or polyphyletic (Tsutsumi & Kato,
2005). Paleotropics and subtropics, Pacific Basin. Char-
acters: plants epiphytic (most genera) or epipetric; rhi-
zomes long-creeping, dictyostelic, dorsiventral, bearing
scales; old leaves cleanly abscising at petiole bases;
blades usually 1–4-pinnate (rarely simple), monomor-
phic (rarely dimorphic); veins free, forking or pinnate;
indument generally lacking on blades and axes, but
sometimes of articulate hairs; sori abaxial, inframarginal
to well back from the margin, ± round, with cup-shaped
to reniform or lunate indusia (rarely forming a submar-
ginal coenosorus in Parasorus); sporangia with 3-rowed,
usually long stalks; annuli vertical; spores ellipsoid,
monolete, yellowish to tan, perine various, but usually
not strongly winged or cristate; gametophytes green, cor-
date; x = 40.
37. Family Polypodiaceae. —
Polygrams; incl.
Drynariaceae, Grammitidaceae (grammitids), Gymno-
grammitidaceae, Loxogrammaceae, Platyceriaceae,
Pleurisoriopsidaceae. Ca. 56 genera, ca. 1200 spp. Pan-
tropical, a few temperate. Genera include Acrosorus,
Adenophorus (Ranker & al., 2003), Aglaomorpha (incl.
Photinopteris, Merinthosorus, Pseudodrynaria, Holosta-
chyum; Janssen & Schneider, 2005), Arthromeris, Belv-
isia, Calymmodon, Campyloneurum, Ceradenia, Christi-
opteris (Kreier & Schneider, unpubl.), Chrysogrammitis,
Cochlidium, Colysis, Ctenopteris, Dicranoglossum, Dic-
tymia, Drynaria (Janssen & Schneider, 2005),
Enterosora, Goniophlebium s.l., Grammitis, Lecan-
opteris (Haufler & al., 2003), Lellingeria, Lemmaphyl-
lum, Lepisorus (incl. Platygyria), Leptochilus, Loxo-
gramme (incl. Anarthropteris, a monotype from New
Zealand; Kreier & Schneider, in press), Melpomene,
Microgramma (incl. Solanopteris), Micropolypodium,
Microsorum, Neocheiropteris (incl. Neolepisorus),
Neurodium, Niphidium, Pecluma, Phlebodium, Phym-
atosorus, Platycerium (Kreier & Schneider, 2006),
Pleopeltis, Polypodioides, Polypodium, Prosaptia,
Pyrrosia (incl. Drymoglossum), Scleroglossum, Sel-
liguea (incl. Crypsinus, Polypodiopteris), Serpocaulon
(Smith & al., in press), Synammia (Schneider & al.,
2006), Terpsichore, Themelium, Thylacopteris (Schnei-
der & al., 2004a), and Zygophlebia
. Additional mono-
typic genera, include Caobangia
, Drymotaenium,
(Wu & al., 2005), Luisma
Pleurosoriopsis, and Podosorus
Polypodiaceae s.s., as often recognized (e.g., by Kra-
mer in Kubitzki, 1990), is paraphyletic, because it ex-
cludes the grammitids, often segregated as Grammitida-
ceae (Ranker & al., 2004; Schneider & al., 2004d). Ge-
neric boundaries need clarification, and, in particular,
Polypodium and Microsorum, two of the largest assem-
blages, are known to be polyphyletic (Schneider & al.,
2004d). Certain previously misplaced genera are now
shown to be nested within Polypodiaceae, e.g., Pleuro-
soriopsis (Hasebe & al., 1995, Schneider & al., 2004d)
and Gymnogrammitis (Schneider & al., 2002b). Polypo-
diaceae contains large wholly Neotropical and wholly
Paleotropical clades (Schneider & al., 2004d).
Mostly epiphytic and epipetric, a few terrestrial;
pantropical. Characters: rhizomes long-creeping to short-
creeping, dictyostelic, bearing scales; petioles cleanly
abscising near their bases or not (most grammitids), leav-
ing short phyllopodia; blades monomorphic or dimor-
phic, mostly simple to pinnatifid or 1-pinnate (uncom-
monly more divided); indument lacking or of hairs and/
or scales on the blade; veins often anastomosing or retic-
ulate, sometimes with included veinlets, or veins free
(most grammitids); indument various, of scales, hairs, or
glands; sori abaxial (rarely marginal), round to oblong or
elliptic, occasionally elongate, or the sporangia acrosti-
choid, sometimes deeply embedded; sori exindusiate,
sometimes covered by caducous scales when young (e.g.,
Lepisorus, Pleopeltis); sporangia with 1–3-rowed, usual-
ly long stalks, frequently with paraphyses on sporangia
or on receptacle; spores hyaline to yellowish, reniform,
and monolete (non-grammitids), or greenish and glo-
bose-tetrahedral, trilete (most grammitids); perine vari-
ous, usually thin, not strongly winged or cristate; mostly
x = 35, 36, 37 (25 and other numbers also known).
Within Polypodiaceae, as defined above, nest the
grammitid ferns (Ranker & al., 2004; Schneider & al.,
2004d). Tryon & Tryon (1982) and Hennipman (1996)
subsumed the grammitids in Polypodiaceae, as we do
here. Grammitids (ca. 20 genera, 600 species, pantropi-
cal) do share a large number of morphological synapo-
morphies: veins free (mostly); scales lacking on blades;
setiform, often dark red-brown hairs on leaves; sporan-
gial stalks 1-rowed; spores green, trilete; gametophytes
ribbon-shaped. Some genera of grammitids have now
been shown to be polyphyletic, e.g., Ctenopteris, Gram-
mitis, Micropolypodium, and Terpsichore, while others
are likely monophyletic, e.g., Ceradenia, Melpomene,
Prosaptia s.l. (Ranker & al., 2004).
The authors thank Ray Cranfill for comments on an early
draft of the manuscript. We also thank Chie Tsutsumi and
Masahiro Kato, for sharing a pre-publication copy of their paper
on Davalliaceae and related genera, and two anonymous review-
ers, for providing helpful comments. James Reveal kindly provid-
ed comments and suggestions on nomenclatural aspects of our
Smith & al. • Fern classification55 (3) • August 2006: 705–731
work, especially information summarized in appendices. Our work
was supported in part by National Science Foundation grants
DEB-9616260 to ARS; DEB-9615533, DEB-0089909, and DEB-
0347840 to KMP, DEB-9707087 to PGW, and DEB-0408077 to
ES; a postdoctoral fellowship from the Swedish Research Council
(2003-2724) to PK; and a German Science Foundation grant
SCHN 758/2-1 to HS.
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Appendix 1. Supra-ordinal names applied to extant ferns. Boldface indicates names adopted in this classification. *
refers to Kenrick & Crane, 1997.
Subclass Blechnidae Doweld Subdivision Ophioglossophytina Doweld
Subclass Cyatheidae Doweld Class Ophioglossopsida Thomé
Class Cyatheopsida Doweld Subclass Osmundidae Doweld
Subclass Dennstaedtiidae Doweld Class Osmundopsida Doweld
Subclass Equisetidae Warming Subclass Plagiogyriideae Doweld
Division Equisetophyta D. H. Scott Subclass Polypodiidae* Cronquist, Takht. & Zimmerm.
Subdivision Equisetophytina Reveal Division Polypodiophyta Cronquist, Takht. & Zimmerm.
Class Equisetopsida* C. Agardh Subdivision Polypodiophytina Reveal
Subclass Filicidae Hook. & Arn., nom. illeg. Class Polypodiopsida Cronquist, Takht. & Zimmerm.
Division Filicophyta J. Mackay, nom. illeg. Subclass Psilotidae Reveal*
Class Filicopsida C. Agardh, nom. illeg. Division Psilotophyta Heintze
Subclass Gleichenidae Doweld Subdivision Psilotophytina Tippo ex Reveal
Class Gleicheniopsida Doweld Class Psilotopsida* D. H. Scott
Class Hymenophyllopsida Doweld Subclass Pterididae Schmakov
Subclass Marattiidae Klinge Subkingdom Pteridobiotina Underw.
Division Marattiophyta Heintze Division Pteridophyta Schimp.
Class Marattiopsida Doweld Superdivision Pteridophytanae Doweld
Subclass Marsileidae Doweld Subdivision Pteridophytina Engl.
Class Marsileopsida Trevis. Class Pteridopsida Ritgen
Class Matoniopsida Doweld Subclass Salviniidae Cronquist, Takht. & Zimmerm. ex Reveal
Subclass Matoniidae Doweld Class Salviniopsida Kamelin & Schmakov
Infradivision Moniliformopses*, nom. illeg. Subclass Schizaeidae Doweld
Subclass Ophioglossidae* Klinge Class Schizaeopsida Doweld
Division Ophioglossophyta Bek.
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
Appendix 3. Familial names applied to extant ferns, and their taxonomic disposition. Family names accepted by us are
in boldface. Synonyms are in italics. Unpublished or otherwise illegitimate names are in quotation marks. An equal sign
(=) is intended to indicate that we regard the first name as a heterotypic synonym of the family name we adopt.
Acrostichaceae Mett. ex A.B. Frank = Pteridaceae Dicksoniaceae M.R. Schomb., nom. cons. over Thyrsopteridaceae;
Actiniopteridaceae Pic. Serm. = Pteridaceae here included in Cyatheales
Adiantaceae Newman, nom. cons. over Parkeriaceae = Pteridaceae Dicranopteridaceae Ching ex Doweld = Gleicheniaceae
Alsophilaceae C. Presl = Cyatheaceae Dictyoxiphiaceae” Ching, nom. nud. = Tectariaceae
Anemiaceae Link; here included in Schizaeales Didymochlaenaceae” Ching, nom. nud. = Dryopteridaceae, tentatively
Angiopteridaceae Fée ex J. Bommer = Marattiaceae Dipteridaceae Seward & E. Dale; here included in Gleicheniales
Anopteraceae Doweld = Pteridaceae Drynariaceae Ching = Polypodiaceae
Antrophyaceae Ching = Pteridaceae Dryopteridaceae Herter, nom. cons. over Peranemataceae; here
Aspidiaceae” Burnett, nom. illeg. = Dryopteridaceae included in Polypodiales
Aspleniaceae Newman; here included in Polypodiales Elaphoglossaceae Pic. Serm. = Dryopteridaceae
Athyriaceae Alston = Woodsiaceae Equisetaceae Michx. ex DC.; here included in Equisetales
Azollaceae Wettst. = Salviniaceae Filicaceae” Juss., nom. illeg.
Blechnaceae Newman; here included in Polypodiales Gleicheniaceae C. Presl; here included in Gleicheniales
Bolbitidaceae Ching = Dryopteridaceae Grammitidaceae Newman [often misspelled Grammitaceae] =
Botrychiaceae Horan. = Ophioglossaceae Polypodiaceae
Ceratopteridaceae Underw. = Parkeriaceae = Pteridaceae Gymnogrammitidaceae Ching (incl. Gymnogrammaceae, spelling
Cheilanthaceae B.K. Nayar = Pteridaceae variant used by some authors) = Polypodiaceae
Cheiropleuriaceae Nakai = Dipteridaceae Helminthostachyaceae Ching = Ophioglossaceae
Christenseniaceae Ching = Marattiaceae Hemionitidaceae Pic. Serm. = Pteridaceae
Cibotiaceae Korall; here included in Cyatheales Hymenophyllaceae Mart.; here included in Hymenophyllales
Cryptogrammaceae Pic. Serm. = Pteridaceae Hymenophyllopsidaceae Pic. Serm. = Cyatheaceae
Culcitaceae Pic. Serm.; here included in Cyatheales Hypodematiaceae Ching = Dryopteridaceae, tentatively
Cyatheaceae Kaulf.; here included in Cyatheales Hypoderriaceae” Ching, nom. nud., used by various authors, incl.
Cystodiaceae J.R. Croft = Lindsaeaceae Dickason (1946) = Tectariaceae
Cystopteridaceae Schmakov Hypolepidaceae Pic. Serm. = Dennstaedtiaceae
Danaeaceae C. Agardh = Marattiaceae Kaulfussiaceae” Campb., nom. illeg. = Marattiaceae
Davalliaceae M.R. Schomb.; here included in Polypodiales Lindsaeaceae C. Presl; here included in Polypodiales
Dennstaedtiaceae Lotsy; here included in Polypodiales Lomariopsidaceae Alston; here included in Polypodiales
Appendix 2. Ordinal and subordinal names applied to extant ferns. In this list, an equal sign (=) indicates that we regard
the first name as a synonym of the one we adopt. * refers to orders adopted by Pichi Sermolli (1977); most of these
names are not validly published. Boldface indicates orders adopted in our classification. A name in quotes indicates
that it is not validly published.
Order “Aspidiales”: used by various authors, e.g., Order Loxomatales Pic. Serm. ex Reveal = Cyatheales
Pichi Sermolli (1958, 1977), nom. illeg. = Polypodiales Order Marattiales Link
Order Aspleniales Pic. Serm. ex Reveal = Polypodiales Order Marsileales Bartl. = Salviniales
Order Athyriales Schmakov = Polypodiales Order Matoniales Pic. Serm. ex Reveal = Gleicheniales
Order Blechnales Pic. Serm. ex Reveal = Polypodiales Order Metaxyales Doweld = Cyatheales
Order Christenseniales Doweld = Marattiales Order Monachosorales Doweld = Polypodiales
Order Cyatheales A.B. Frank Order Negripteridales Pic. Serm. ex Reveal = Polypodiales
Suborder Cyatheineae Bessey Order Ophioglossales Link
Order “Davalliales”, nom. illeg. = Polypodiales Order Osmundales Link
Order Dennstaedtiales Doweld (name also used by Pichi Suborder Osmundineae Bessey
Sermolli, 1977, where it is a nom. nud.) = Polypodiales Order Parkeriales A.B. Frank = Polypodiales
Order Dicksoniales Pic. Serm. ex Reveal = Cyatheales Order Pilulariales Bercht. & J. Presl = Salviniales
Order Dipteridales Doweld = Gleicheniales Order Plagiogyriales Pic. Serm. ex Reveal = Cyatheales
Order Dryopteridales Schmakov = Polypodiales Order Platyzomatales* Pic. Serm. ex Reveal = Polypodiales
Order Equisetales DC. ex Bercht. & J. Presl Order Polypodiales Link
Suborder Equisetineae Rabenh. Suborder Polypodiineae Bessey
Order Filicales Dumort., nom. illeg.Order Psilotales Prantl
Order Gleicheniales* Link Suborder Psilotineae Engl.
Suborder Gleicheniineae Bessey Order Pteridales* Doweld = Polypodiales
Order “Hydropteridales” (Hydropterides) used by many authors, Order Saccolomatales Doweld = Polypodiales
e.g., Copeland (1947), Kubitzki (1990) = Salviniales Order Salviniales Bartl.
Order Hymenophyllales A.B. Frank Order Schizaeales Schimp.
Suborder Hymenophyllineae Bessey Suborder Schizaeineae Bessey
Order Hymenophyllopsidales Pic. Serm. ex Reveal = Cyatheales Order Stromatopteridales* Pic. Serm. ex Reveal = Gleicheniales
Order Lindsaeales Doweld = Polypodiales Order Thelypteridales Doweld = Polypodiales
Order Lonchitidales Doweld = Polypodiales
Smith & al. • Fern classification55 (3) • August 2006: 705–731
Appendix 4. Familial, ordinal, and supra-ordinal names for extant ferns, with citations from Reveal & Doweld (subm.),
as revised from Hoogland & Reveal (2005), to comply with the Vienna Code. Orthographic variations in spelling, in orig-
inal references, are in quotation marks. Italicized names are not validly published. * refers to ordinal names used by
Pichi Sermolli (1977), some of them unpublished, some published by earlier authorities. Commonly used “descriptive
names” (Art. 16.1), e.g., Hydropterides, are also given, but no attempt has been made to include all such names, or to
determine first usage; no good index is available for these.
Fam. Acrostichaceae Mett. ex A.B. Frank, Syn. Pflanzenk., ed. 2, 3: 1453, 1458. 1874; nom. rej. in favor of Adiantaceae.
Fam. Actiniopteridaceae Pic. Serm., Webbia 17: 5. 31 Aug 1962.
Fam. Adiantaceae Newman, Hist. Brit. Ferns: 5. 1–5. Feb 1840; nom. cons. over Parkeriaceae (1825).
Fam. Alsophilaceae C. Presl, Gefässbündel Farrn: 22 (footnote), 32. 1847.
Fam. Anemiaceae Link, Fil. Spec.: 23. 3–10 Sep 1841.
Fam. Angiopteridaceae Fée ex J. Bommer, Bull. Soc. Bot. Belgique 5: 345, 359. before 28 May 1867 (“Angiopterideae”).
Fam. Anopteraceae Doweld, Tent. Syst. Pl. Vasc.: li. 23 Dec 2001.
Fam. Antrophyaceae Ching, Acta Phytotax. Sin. 16(3): 11. Aug 1978. – basionym: Antrophyoideae Link, Fil. Spec.: 140. 3–10 Sep 1841
Fam. Aspidiaceae Burnett, Outl. Bot.: 324, 1156. Feb 1835; nom. illeg. (Arts. 18.3). The name is illegitimate because it was based on an
illegitimate generic name (Art. 18.3).
Order Aspidiales*, nom. illeg., used by various authors, e.g., Pichi Sermolli (1958, 1977); does not satisfy Art. 16.1(a), that a name be taken
from a legitimate name of an included family.
Fam. Aspleniaceae Newman, Hist. Brit. Ferns: 6. 1–5 Feb 1840.
Order Aspleniales Pic. Serm. ex Reveal, Phytologia 79: 72. 29 Apr 1996.
Fam. Athyriaceae Alston, Taxon 5: 25. 25 Mar 1956.
Order Athyriales Schmakov, Turczaninowia 4: 55. 15 May 2001.
Fam. Azollaceae Wettst., Handb. Syst. Bot. 2: 77. Nov 1903.
Fam. Blechnaceae Newman, Hist. Brit. Ferns, ed. 2: 8. 1844.
Order Blechnales* Pic. Serm. ex Reveal, Phytologia 74: 175. 25 Mar 1993.
Subclass Blechnidae Doweld, New Syllabus: 368. May 2006.
Fam. Bolbitidaceae Ching, Acta Phytotax. Sin. 16(3): 15. Aug 1978. – basionym: Bolbitideae Pic. Serm., Webbia 23: 381. 30 Jun 1969.
Fam. Botrychiaceae Horan., Char. Ess. Fam.: 15. 30 Jun 1847.
Fam. Ceratopteridaceae Underw., Our Native Ferns, ed. 6: 65, 78. Jul–Aug 1900.
Fam. Cheilanthaceae B.K. Nayar, Taxon 19: 233. 29 Apr 1970.
Fam. Cheiropleuriaceae Nakai, Bot. Mag. (Tokyo) 42: 210. Apr 1928.
Fam. Christenseniaceae Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 227. 25 Dec 1940.
Order Christenseniales Doweld, Tent. Syst. Pl. Vasc.: vii. 23 Dec 2001.
Fam. Cibotiaceae Korall in A. R. Smith & al., Taxon 55: 712. 2006. – basionym: Cibotioideae B. K. Nayar, Taxon 19: 234. 1970.
Fam. Cryptogrammaceae Pic. Serm., Webbia 17: 299. 20 Apr 1963.
Fam. Culcitaceae Pic. Serm., Webbia 24: 702. 28 Apr 1970.
Fam. Cyatheaceae Kaulf., Wesen Farrenkr.: [119]. Jul–Sep 1827.
Order Cyatheales A.B. Frank, Syn. Pflanzenk., ed. 2, 3: 1452, 1456. 1877 (“Cyatheaceae”).
Subclass Cyatheidae Doweld, Tent. Syst. Pl. Vasc.: xii. 23 Dec 2001.
Suborder Cyatheineae Bessey, Bot. High Schools: 376. 14–21 Aug 1880 (“Cyatheaceae”).
Class Cyatheopsida Doweld, Tent. Syst. Pl. Vasc.: xii. 23 Dec 2001.
Fam. Cystodiaceae J.R. Croft, Kew Bull. 41: 797. 20 Oct 1986.
Appendix 3. Continued.
Lonchitidaceae Doweld = Lindsaeaceae Platyceriaceae Ching = Polypodiaceae
Lophosoriaceae Pic. Serm.; here included in Dicksoniaceae Platyzomataceae Nakai = Pteridaceae
Loxogrammaceae Ching ex Pic. Serm. = Polypodiaceae Pleurosoriopsidaceae Kurita & Ikebe ex Ching = Polypodiaceae
Loxomataceae C. Presl [often misspelled “Loxsomaceae”]; here Polypodiaceae J. Presl; here included in Polypodiales
included in Cyatheales Psilotaceae J.W. Griff. & Henfr.; here included in Psilotales
Lygodiaceae M. Roem.; here included in Schizaeales Pteridaceae E.D.M. Kirchn.; here included in Polypodiales
Marattiaceae Kaulf., nom. cons. prop.; here included in Marat- Pteridiaceae Ching = Dennstaedtiaceae
tiales; antedated by Danaeaceae (Murdock & al., subm.) Saccolomataceae Doweld; here included in Polypodiales
Marsileaceae Mirb.; here included in Salviniales Salviniaceae Martynov; here included in Salviniales
Matoniaceae C. Presl; here included in Gleicheniales Schizaeaceae Kaulf.; here included in Schizaeales
Metaxyaceae Pic. Serm.; here included in Cyatheales Sinopteridaceae Koidz., nom. rej. in favor of Adiantaceae =
Mohriaceae C.F. Reed. = Anemiaceae Pteridaceae
Monachosoraceae Ching = Dennstaedtiaceae Sphaerostephanaceae” Ching, nom. nud. = Thelypteridaceae
Negripteridaceae Pic. Serm. = Pteridaceae Stenochlaenaceae Ching = Blechnaceae
Nephrolepidaceae Pic. Serm. = Lomariopsidaceae, tentatively Stromatopteridaceae Bierh. = Gleicheniaceae
Oleandraceae Ching ex Pic. Serm.; here included in Polypodiales Taenitidaceae Pic. Serm. = Pteridaceae
Onocleaceae Pic. Serm.; here included in Polypodiales Tectariaceae Panigrahi; here included in Polypodiales
Ophioglossaceae Martynov; here included in Ophioglossales Thelypteridaceae Pic. Serm.; here included in Polypodiales
Osmundaceae Martynov; here included in Osmundales Thyrsopteridaceae C. Presl; here included in Cyatheales
Parkeriaceae Hook. = Pteridaceae Tmesipteridaceae Nakai = Psilotaceae
Peranemataceae (C. Presl) Ching = Dryopteridaceae Trichomanaceae Burmeist. = Hymenophyllaceae
Pilulariaceae Mirb. ex DC. (Pilulariae) = Marsileaceae Vittariaceae Ching = Pteridaceae
Plagiogyriaceae Bower; here included in Cyatheales Woodsiaceae Herter: here included in Polypodiales
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
Appendix 4. Continued.
Fam. Cystopteridaceae Schmakov, Turczaninowia 4: 60. 15 May 2001. – basionym: Cystopteridinae Payer, Bot. Crypt.: 198. 1850
Fam. Danaeaceae C. Agardh, Aphor. Bot. 117. 19 Jun 1822; nom. rej. prop. in favor of Marattiaceae, nom. cons. prop. (Murdock & al., subm.).
Fam. Davalliaceae M.R. Schomb., Reis. Br.-Guiana 2: 883. 12–15 Jul 1848. – basionym: Davallieae Gaudich. in Freyc., Voy. Uraniae, Bot. 370.
18 Jul 1829.
Order Davalliales, nom. illeg.: used by Pichi Sermolli (1958).
Fam. Dennstaedtiaceae Lotsy, Vortr. Bot. Stammesgesch. 2: 655. 9 Aug 1909 (“Dennstaedtineaceae”).
Order Dennstaedtiales Doweld, Tent. Syst. Pl. Vasc.: x. 23 Dec 2001.
Subclass Dennstaedtiidae Doweld, Tent. Syst. Pl. Vasc.: x. 23 Dec 2001.
Fam. Dicksoniaceae M.R. Schomb., Reis. Br.-Guiana 2: 1047. 12–15 Jul 1848; nom. cons. over Thyrsopteridaceae (1847) – basionym:
Dicksonieae Gaudich., in Freycinet, Voy. Uranie, Bot.: 367. 18 Jul 1829.
Order Dicksoniales* Pic. Serm. ex Reveal, Phytologia 74: 175. 25 Mar 1993.
Fam. Dicranopteridaceae Ching ex Doweld, Tent. Syst. Pl. Vasc.: x. 23 Dec 2001; nom. invalid. in Ching, Acta Phytotax. Sin. 3: 94. 1954.
Fam. Dictyoxiphiaceae Ching, Sunyatsenia 5: 218. 1940; nom. nud., without Latin description.
Fam. Didymochlaenaceae Ching, Sunyatsenia 5: 253. 1940; nom. nud., without Latin description.
Fam. Dipteridaceae Seward & E. Dale, Philos. Trans., ser. B, 194: 487, 499, 502. Dec 1901 (“Dipteridinae”). – basionym: Dipteridinae Diels, in
Engler & Prantl, Nat. Pflanzenfam. I, 4: 167. 21 Jul 1899.
Order Dipteridales Doweld, Tent. Syst. Pl. Vasc.: x. 23 Dec 2001.
Fam. Drynariaceae Ching, Acta Phytotax. Sin. 16(4): 19. Nov 1978.
Fam. Dryopteridaceae Herter, Rev. Sudamer. Bot. 9: 15. 1949; nom. cons. over Peranemataceae Ching.
Order Dryopteridales Schmakov, Turczaninowia 4: 66, 67. 15 May 2001.
Fam. Elaphoglossaceae Pic. Serm., Webbia 23: 209. 20 Nov 1968.
Fam. Equisetaceae Michx. ex DC., Essai Propr. Méd. Pl.: 49. Jul 1804.
Order Equisetales DC. ex Bercht. & J. Presl, Prir. Rostlin: 271. Jan–Apr 1820 (“Equisetaceae”).
Subclass Equisetidae Warming, Osnov. Bot.: 221. 22–28 Apr 1883 (“Equisetinae”).
Suborder Equisetineae Rabenh., Deutschl. Krypt.-Fl. 2: xvi, 332. Jan–Feb 1849 (“Equisetaceae”).
Division Equisetophyta D.H. Scott, Stud. Foss. Pl.: 13, 489, 492, 493. Sep 1900 (“Equisetales”).
Subdivision Equisetophytina Reveal, Phytologia 79: 70. 29 Apr 1996.
Class Equisetopsida C. Agardh, Classes Pl.: 7. 20 May 1825 (“Equisetaceae”).
Fam. Filicaceae Juss., Gen. Pl.: 14. 4 Aug 1789. (“Filices”); nom. illeg. (Art. 18.3).
Order Filicales Dumort., Anal. Fam. Pl.: 67. 1829. (“Filicarieae”); nom. illeg. (Art. 16.1).
Subclass Filicidae Hook. & Arn., Brit. Fl., ed. 6, 1: 564. Jul 1850. (“Filices”); nom. illeg. (Art. 16.1).
Division Filicophyta J. Mackay, Fl. Hiber.: 336. Jul 1836; nom. illeg. (Art. 16.1).
Class Filicopsida C. Agardh, Aphor. Bot.: 114. 19 Jul 1822. (“Filices”); nom. illeg. (Art. 16.1). Used by many authors, including K&C, 1997, for
what we here call Polypodiopsida.
Fam. Gleicheniaceae C. Presl, Reliq. Haenk. 1: 70. Jun–Nov 1825 (“Gleicheniae”). – basionym: [unranked] Gleicheneae R. Br., Prodr. 160. 27
Mar 1810.
Order Gleicheniales* Link, Handbuch 3: 10. 1833 (“Gleicheneae”).
Subclass Gleichenidae Doweld, New Syllabus: 337. May 2006.
Suborder Gleicheniineae Bessey, Bot. High Schools: 376. 14–21 Aug 1880 (“Gleicheniaceae”).
Class Gleicheniopsida Doweld, Tent. Syst. Pl. Vasc.: ix. 23 Dec 2001.
Fam. Grammitidaceae Newman, Hist. Brit. Ferns: 7. 1–5 Feb 1840 (“Grammitideae”). Often misspelled Grammitaceae; see, e.g., Pichi Sermolli,
Fam. Gymnogrammitidaceae Ching, Acta Phytotax. Sin. 11: 12. Jan 1966. Often misspelled Gymnogrammaceae.
Fam. Helminthostachyaceae Ching, Bull. Fan Mem. Inst. Biol., Bot. 10: 235. 15 Mar 1941.
Fam. Hemionitidaceae Pic. Serm., Webbia 21: 487. 15 Jul 1966.
Order Hydropterides Campbell, Mosses and Ferns: 304, 307. 1895. Descriptive name (Art. 16.1).
Fam. Hymenophyllaceae Mart., Consp. Regni Veg.: 3. Sep–Oct 1835.
Order Hymenophyllales* A.B. Frank, Syn. Pflanzenk., ed. 2, 3: 1452, 1474. 1877 (“Hymenophyllaceae”).
Suborder Hymenophyllineae Bessey, Bot. High Schools: 376. 14–21 Aug 1880 (“Hymenophyllaceae”).
Class Hymenophyllopsida Doweld, Tent. Syst. Pl. Vasc.: ix. 23 Dec 2001.
Fam. Hymenophyllopsidaceae Pic. Serm., Webbia 24: 712. 28 Apr 1970.
Order Hymenophyllopsidales Pic. Serm. ex Reveal, Phytologia 74: 175. 25 Mar 1993.
Fam. Hypodematiaceae Ching, Acta Phytotax. Sin. 13(1): 96. Jan 1975.
Fam. Hypoderriaceae Ching, Sunyatsenia 5: 245. 1940; nom. nud., without Latin description.
Fam. Hypolepidaceae Pic. Serm., Webbia 24: 705. 28 Apr 1970.
Fam. Kaulfussiaceae Campb., Evol. Land Pl. 333. 1940; nom. illeg. (Art. 18.3), because it is based on the illegitimate generic name Kaulfussia
Blume (1828), non Dennstedt (1818), nec Nees (1820).– basionym: Kaulfussioideae Campb., Struct. Moss. Ferns, ed. 2, 298, 300. Sep 1905
(“Kaulfussieae”), nom. illeg. See also Pichi Sermolli Webbia 12: 353. 1957.
Fam. Lindsaeaceae C. Presl, in M. R. Schomb., Reis. Br.-Guiana 2: 883. 12–15 Jul 1848.
Order Lindsaeales Doweld, New Syllabus: 353. May 2006.
Fam. Lomariopsidaceae Alston, Taxon 5: 25. 25 Mar 1956.
Fam. Lonchitidaceae Doweld, New Syllabus: 353. May 2006.
Order Lonchitidales Doweld, New Syllabus: 353. May 2006.
Fam. Lophosoriaceae Pic. Serm., Webbia 24: 700. 28 Apr 1970.
Fam. Loxogrammaceae Ching ex Pic. Serm., Webbia 29: 11. 10 Feb 1975.
Fam. Loxomataceae C. Presl, Gefässbündel Farrn: 31. 1847 (“Loxsomaceae”).
Order Loxomatales* Pic. Serm. ex Reveal, Phytologia 74: 175. 25 Mar 1993 (“Loxsomales”).
Fam. Lygodiaceae M. Roem., Handb. Allg. Bot. 3: 520. 1840 (“Lygodieae”).
Fam. Marattiaceae Kaulf., Enum. Filic.: 31. 8 Apr–29 May 1824; nom. cons. prop. over Danaeaceae (1822) (Murdock & al., subm.).
Smith & al. • Fern classification55 (3) • August 2006: 705–731
Appendix 4. Continued.
Order Marattiales Link, Hort. Berol. 2: 148. Jul–Dec 1833 (“Marattiaceae”).
Subclass Marattiidae Klinge, Fl. Est-Liv-Churland 1: 93. 22–28 Jun 1882 (“Marattiaceae”).
Division Marattiophyta Heintze, Cormofyt. Fylog.: 22. 1927.
Class Marattiopsida Doweld, Tent. Syst. Pl. Vasc.: vii. 23 Dec 2001.
Fam. Marsileaceae Mirb. in Lam. & Mirb., Hist. Nat. Vég. 5: 126. 21 Nov 1802 (“Marsileae”).
Order Marsileales Bartl., in Martius, Consp. Regn. Veg.: 4. Sep–Oct 1835 (“Marsileaceae”).
Subclass Marsileidae Doweld, Tent. Syst. Pl. Vasc.: x. 23 Dec 2001.
Class Marsileopsida Trevis., Bull. Soc. Roy. Bot. Belgique 16: 5. after 6 May 1877 (“Marsigliales”).
Fam. Matoniaceae C. Presl, Gefässbündel Farrn: 32. 1847.
Order Matoniales* Pic. Serm. ex Reveal, Phytologia 74: 175. 25 Mar 1993.
Subclass Matoniidae Doweld, New Syllabus: 338. May 2006.
Class Matoniopsida Doweld, Tent. Syst. Pl. Vasc.: vii. 23 Dec 2001.
Fam. Metaxyaceae Pic. Serm., Webbia 24: 701. 28 Apr 1970.
Order Metaxyales Doweld, Tent. Syst. Pl. Vasc.: xii. 23 Dec 2001.
Fam. Mohriaceae C.F. Reed, Bol. Soc. Brot., ser. 2, 21: 168. 11 May 1948.
Fam. Monachosoraceae Ching, Acta Phytotax. Sin. 16(4): 17. Nov 1978.
Order Monachosorales Doweld, New Syllabus: 356. May 2006.
Infradivision Moniliformopses Kenrick & P. Crane; nom. nud. in Kenrick & Crane, 1997.
Fam. Negripteridaceae Pic. Serm., Nuovo Giorn. Bot. Ital., ser. 2, 53: 160. 25 Nov 1946.
Order Negripteridales* Pic. Serm. ex Reveal, Phytologia 74: 176. 25 Mar 1993.
Fam. Nephrolepidaceae Pic. Serm., Webbia 29: 8. 10 Feb 1975.
Fam. Oleandraceae Ching ex Pic. Serm., Webbia 20: 745. 20 Dec 1965.
Fam. Onocleaceae Pic. Serm., Webbia 24: 708. 28 Apr 1970.
Fam. Ophioglossaceae Martynov, Tekhno-Bot. Slovar: 438. 3 Aug 1820 (“Ophioglosseae”) – basionym: [unranked] Ophioglosseae R. Br., Prodr.:
163. 27 Mar 1810.
Order Ophioglossales Link, Hort. Berol. 2: 151. Jul–Dec 1833 (“Ophioglossaceae”).
Subclass Ophioglossidae Klinge, Fl. Est-Liv-Churland 1: 94. 22–28 Jun 1882 (“Ophioglosseae”).
Division Ophioglossophyta Bek., Kurs Bot. 1: 89. 1863 (“Ophioglosseae”).
Subdivision Ophioglossophytina Doweld, Tent. Syst. Pl. Vasc.: ix. 23 Dec 2001.
Class Ophioglossopsida Thomé, Lehrb. Bot., ed. 4: 205. 31 Mar 1874 (“Ophioglosseae”).
Fam. Osmundaceae Martynov, Tekhno-Bot. Slovar: 445. 3 Aug 1820.
Order Osmundales Link, Hort. Berol. 2: 138. Jul–Dec 1833 (“Osmundaceae”).
Subclass Osmundidae Doweld, New Syllabus: 334. May 2006.
Suborder Osmundineae Bessey, Bot. High Schools: 377. 14–21 Aug 1880 (“Osmundaceae”).
Class Osmundopsida Doweld, Tent. Syst. Pl. Vasc.: ix. 23 Dec 2001.
Fam. Parkeriaceae Hook., Exot. Fl. 2: ad t. 147. Mar 1825; nom. rej. in favor of Adiantaceae.
Order Parkeriales A.B. Frank, Syn. Pflanzenk., ed. 2, 3: 1452, 1458. 1877 (“Parkeriaceae”).
Fam. Peranemataceae Ching, Sunyatsenia 5: 208, 246. 30 Oct 1940 (“Perenemaceae”); nom. rej. in favor of Dryopteridaceae Herter (1949). –
basionym: Peranemateae C. Presl, [Tent. pterid.] Abh. Königl. Böhm. Ges. Wiss., ser. 4, 5. 64. before 2 Dec 1836.
Fam. Pilulariaceae Mirb. ex DC., Essai Propr. Méd. Pl.: 48. Jul 1804 (“Pilulariae”).
Order Pilulariales Bercht. & J. Presl, Prir. Rostlin: 272. Jan–Apr 1820 (“Pilulariae”).
Fam. Plagiogyriaceae Bower, Ann. Bot. (London) 40: 484. Apr 1926.
Order Plagiogyriales Pic. Serm. ex Reveal, Phytologia 74: 176. 25 Mar 1993.
Subclass Plagiogyriidae Doweld, Tent. Syst. Pl. Vasc.: xii. 23 Dec 2001.
Fam. Platyceriaceae Ching, Acta Phytotax. Sin. 16(3): 18. Aug 1978.
Fam. Platyzomataceae Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 4. Dec 1950.
Order Platyzomatales* Pic. Serm. ex Reveal, Phytologia 74: 176. 25 Mar 1993.
Fam. Pleurosoriopsidaceae Kurita & Ikebe ex Ching, Acta Phytotax. Sin. 16(4): 17. Nov 1978.
Fam. Polypodiaceae J. Presl & C. Presl, Delic. Prag.: 159. Jul 1822.
Order Polypodiales* Link, Hort. Berol. 2: 5. Jul–Dec 1833 (“Polypodiaceae”).
Subclass Polypodiidae Cronquist, Takht. & Zimmerm., Taxon 15: 133. Apr 1966.
Suborder Polypodiineae Bessey, Bot. High Schools: 377. 14–21 Aug 1880 (“Polypodiaceae”).
Division Polypodiophyta Cronquist, Takht. & Zimmerm., Taxon 15: 133. Apr 1966.
Subdivision Polypodiophytina Reveal, Phytologia 79: 70. 29 Apr 1996.
Class Polypodiopsida Cronquist, Takht. & Zimmerm., Taxon 15: 133. Apr 1966.
Fam. Psilotaceae J.W. Griff. & Henfr., Microgr. Dict.: 540. 1 Nov 1855 (“Psiloteae”).
Order Psilotales Prantl, Lehrb. Bot., ed. 5: 183. 5 Jan 1884 (“Psilotaceae”).
Subclass Psilotidae Reveal, Phytologia 79: 70. 29 Apr 1996.
Suborder Psilotineae Engl., Syllabus, ed. 2: 64. May 1898.
Division Psilotophyta Heintze, Cormofyt. Fylog.: 22. 1927.
Subdivision Psilotophytina Tippo ex Reveal, Phytologia 79: 70. 29 Apr 1996.
Class Psilotopsida D.H. Scott, Stud. Foss. Pl., ed. 2: 616, 631, 632. May 1909.
Fam. Pteridaceae E.D.M. Kirchn., Schul-Bot. 109. 13–20 Oct 1831 (“Pteroideae”). Often cited as Pteridaceae Reichb., Handb. Nat.
Pflanzensyst. 138. 1837.
Order Pteridales* Doweld, Tent. Syst. Pl. Vasc.: xi. 23 Dec 2001.
Fam. Pteridiaceae Ching, Acta Phytotax. Sin. 13(1): 96. Jan 1975.
Subclass Pterididae Schmakov, Turczaninowia 4: 42. 15 May 2001.
Subkingdom Pteridobiotina Underw., in Britton & A. Br., Illus. Fl. N. U.S. 1: 1. 15 Aug 1896 (“Pteridophyta”).
Division Pteridophyta Schimp. in Zittel, Handb. Palaeontol. 2(1): 1. 15 Sep 1879.
Superdivision Pteridophytanae Doweld, Tent. Syst. Pl. Vasc.: v. 23 Dec 2001.
Smith & al. • Fern classification 55 (3) • August 2006: 705–731
Appendix 5. Index to genera with family assignments proposed in this classification. All accepted genera (but not all
synonyms) in Kramer in Kubitzki (1990) are accounted for here. Genera newly described or recircumscribed since 1990
are also included. Accepted names in roman; synonyms in italics. Family numbers assigned in text.
Abacopteris = Cyclosorus Archangiopteris = Angiopteris Ceterach = Asplenium Cyathea (20)
Abrodictyum (6) Argyrochosma (26) Ceterachopsis = Asplenium Cyclodium (32)
Acrophorus (32) Arthromeris (37) Cheilanthes (26) Cyclogramma = Cyclosorus
Acrorumohra (32) Arthropteris (34) Cheilanthopsis (29) Cyclopeltis (33)
Acrosorus (37) Aspidotis (26) Cheiloplecton (26) Cyclosorus (28)
Acrostichum (26) Aspleniopsis = Austrogramme Cheiroglossa = Ophioglossum Cyrtogonellum (32)
Actiniopteris (26) Asplenium (27) Cheiropleuria (8) Cyrtomidictyum (32)
Actinostachys (12) Astrolepis (26) Chieniopteris = Woodwardia Cyrtomium (32)
Acystopteris (29) Ataxipteris (32) Chingia = Cyclosorus Cystodium (23)
Adenoderris (32) Athyriopsis = Deparia Chlamydogramme = Tectaria Cystopteris (29)
Adenophorus (37) Athyrium (29) Christella = Cyclosorus Danaea (4)
Adiantopsis (26) Austrogramme (26) Christensenia (4) Davallia (36)
Adiantum (26) Azolla (14) Christiopteris (37) Davalliopsis = Trichomanes
Aenigmopteris (34) Belvisia (37) Chrysochosma = Notholaena Davallodes (36)
Afropteris = Pteris Blechnum (30) Chrysogrammitis (37) Dennstaedtia (25)
Aglaomorpha (37) Blotiella (25) Cibotium (19) Deparia (29)
Aleuritopteris (26) Bolbitis (32) Cionidium = Tectaria Diacalpe = Peranema
Allantodia = Diplazium Bommeria (26) Cnemidaria = Cyathea Dicksonia (21)
Alsophila (20) Botrychium (1) Cochlidium (37) Dicranoglossum (37)
Amauropelta = Thelypteris Botrypus = Botrychium Colysis (37) Dicranopteris (7)
Ampelopteris = Cyclosorus Brainea (30) Coniogramme (26) Dictymia (37)
Amphiblestra = Tectaria Callipteris = Diplazium Coptodipteris (25) Dictyocline = Cyclosorus
Amphineuron = Cyclosorus Callistopteris (6) Cornopteris (29) Dictyodroma = Deparia
Ananthacorus (26) Calochlaena (21) Coryphopteris = Thelypteris Dictyoxiphium = Tectaria
Anarthropteris = Loxogramme Calymmodon (37) Cosentinia (26) Didymochlaena (32)
Anchistea = Woodwardia Camptodium = Tectaria Costaricia = Dennstaedtia Didymoglossum (6)
Anemia (11) Camptosorus = Asplenium Coveniella (32) Diellia = Asplenium
Anetium (26) Campyloneurum (37) Crepidomanes (6) Diplaziopsis (29)
Angiopteris (4) Caobangia (37) Crypsinus = Selliguea Diplazium (29)
Anogramma (26) Cardiomanes = Hymenophyllum Cryptogramma (26) Diplopterygium (7)
Anopteris = Pteris Cassebeera (26) Ctenitis (32) Dipteris (8)
Antigramma = Asplenium Cephalomanes (6) Ctenitopsis = Tectaria Doodia (30)
Antrophyum (26) Ceradenia (37) Ctenopteris (37) Doryopteris (26)
Arachniodes (32) Ceratopteris (26) Culcita (17) Drymoglossum = Pyrrosia
Araiostegia (36) Cerosora (26) Currania = Gymnocarpium Drymotaenium (37)
Appendix 4. Continued.
Subdivision Pteridophytina Engl., Führer Garten Breslau: 10, 11. May 1886 (“Pteridophyta”). Class Pteridopsida Ritgen,
Aufeinanderfolge Org. Gest.: 63. 1828 (“Filices s. Pterides”).
Fam. Saccolomataceae Doweld, New Syllabus: 354. May 2006.
Order Saccolomatales Doweld, New Syllabus: 354. May 2006.
Fam. Salviniaceae Martynov, Tekhno-Bot. Slovar: 559. 3 Aug 1820 (“Salviniae”).
Order Salviniales Link, Hort. Berol. 3: 155. Jul–Dec 1833 (“Salviniaceae”).
Subclass Salviniidae Pic. Serm. ex Reveal, Phytologia 79: 70. 29 Apr 1996.
Class Salviniopsida Kamelin & Schmakov, in Kamelin, Fl. Altaia: 253. 15 May 2005.
Fam. Schizaeaceae Kaulf., Wesen Farrenkr.: [119]. Jul–Sep 1827.
Order Schizaeales* Schimp., Traité Paléont. Vég. 1: 674. Mar 1869.
Subclass Schizaeidae Doweld, New Syllabus: 346. May 2006.
Suborder Schizaeineae Bessey, Bot. High Schools: 377. 14–21 Aug 1880 (“Schizaeaceae”).
Class Schizaeopsida Doweld, Tent. Syst. Pl. Vasc.: x. 23 Dec 2001.
Fam. Sinopteridaceae Koidz., Acta Phytotax. Geobot. 3: 50. 30 May–20 Jul 1934.
Fam. Sphaerostephanaceae Ching, Sunyatsenia 5: 240. 1940; nom. nud., without Latin description.
Fam. Stenochlaenaceae Ching, Acta Phytotax. Sin. 16(4): 18. Nov 1978.
Fam. Stromatopteridaceae Bierh., Phytomorphology 18: 263. 15 Dec 1968. – basionym: Stromatopteridoideae Nakai, Bull. Natl. Sci.
Mus. Tokyo 29: 32. 1950.
Order Stromatopteridales* Pic. Serm. ex Reveal, Phytologia 74: 176. 25 Mar 1993.
Fam. Taenitidaceae Pic. Serm., Webbia 29: 1. 10 Feb 1975.
Fam. Tectariaceae Panigrahi, J. Orissa Bot. Soc. 8: 41. 1986.
Fam. Thelypteridaceae Pic. Serm., Webbia 24: 709. 28 Apr 1970.
Order Thelypteridales Doweld, Tent. Syst. Pl. Vasc.: xi. 23 Dec 2001.
Fam. Thyrsopteridaceae C. Presl, Gefässbündel Farrn: 22, 38. 1847 (“Thyrsopterideae”); nom. rej. in favor of Dicksoniaceae (1848).
Fam. Tmesipteridaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.] 206. 1943.
Fam. Trichomanaceae Burmeist., Handb. Naturgesch. 1: 196. 12–17 Dec 1836 (“Trichomanoideae”).
Fam. Vittariaceae Ching, Sunyatsenia 5: 210, 232. 30 Oct 1940. – basionym: Vittarieae C. Presl, [Tent. Pterid.] Abh. Königl. Böhm.
Ges. Wiss., ser. 4, 5: 164. before 2 Dec 1836 (“Vittariaceae”).
Fam. Woodsiaceae Herter, Revista Sudamer. Bot. 9: 14. Jun 1949. – basionym: Woodsieae A. Gray, Man. Bot., ed. 2. 588. 1 Sep 1856.
Smith & al. • Fern classification55 (3) • August 2006: 705–731
Appendix 5. Continued.
Drynaria (37) Leucostegia (32) Ophioderma = Ophioglossum Pyrrosia (37)
Dryoathyrium = Deparia Lindsaea (23) Ophioglossum (1) Quercifilix = Tectaria
Dryopolystichum (32) Lindsayoides = Nephrolepis Oreopteris = Thelypteris Radiovittaria (26)
Dryopsis (32) Lithostegia (32) Ormoloma (23) Regnellidium (13)
Dryopteris (32) Litobrochia = Pteris Orthiopteris = Saccoloma Revwattsia (32)
Edanyoa = Bolbitis Llavea (26) Osmunda (5) Rhachidosorus (29)
Egenolfia = Bolbitis Lomagramma (32) Pachypleuria (36) Rheopteris (26)
Elaphoglossum (32) Lomaphlebia = Grammitis? Paesia (25) Rosenstockia = Hymenophyllum
Enterosora (37) Lomaria = Blechnum Paltonium = Neurodium Rumohra (32)
Equisetum (3) Lomariopsis (33) Papuapteris = Polystichum Saccoloma (24)
Eriosorus (26) Lonchitis (23) Paraceterach (26) Sadleria (30)
Fadyenia = Tectaria Lophosoria (21) Parahemionitis (26) Saffordia = Trachypteris
Feea = Trichomanes Lorinseria = Woodwardia Parasorus = Davallia Sagenia = Tectaria
Fourniera = Sphaeropteris Loxogramme (37) Parathelypteris = Thelypteris Salpichlaena (30)
Glaphyropteridopsis = Cyclosorus Loxoma (16) Pecluma (37) Salvinia (14)
Glaphyropteris = Cyclosorus Loxoscaphe = Asplenium Pelazoneuron = Cyclosorus Sceptridium = Botrychium
Gleichenella (7) Loxsomopsis (16) Pellaea (26) Schaffneria = Asplenium
Gleichenia (7) Luisma (37) Peltapteris = Elaphoglossum Schizaea (12)
Goniophlebium (37) Lunathyrium = Deparia Pentagramma (26) Scleroglossum (26)
Goniopteris = Cyclosorus Lygodium (10) Pentarhizidium (31) Scoliosorus (26)
Gonocormus = Crepidomanes Macroglena = Abrodictyum Peranema (32) Scyphularia = Davallia
Grammitis (37) Macrothelypteris (28) Phanerophlebia (32) Selliguea (37)
Gymnocarpium (29) Mankyua (1) Phanerosorus (9) Serpocaulon (37)
Gymnogramma = Hemionitis Marattia (4) Phegopteris (28) Serpyllopsis = Hymenophyllum
Gymnogrammitis (37) Marginariopsis = Pleopeltis Phlebodium (37) Sinephropteris = Asplenium
Gymnopteris = Hemionitis Marsilea (13) Photinopteris = Aglaomorpha Sinopteris = Aleuritopteris
Gymnosphaera = Alsophila Matonia (9) Phyllitis = Asplenium Solanopteris = Microgramma
Haplopteris (26) Matteuccia (31) Phymatosorus (37) Sphaerocionium =
Hecistopteris (26) Maxonia (32) Pilularia (13) Hymenophyllum
Helminthostachys (1) Mecodium = Hymenophyllum Pityrogramma (26) Sphaeropteris (20)
Hemidictyum (29) Megalastrum (32) Plagiogyria (18) Sphaerostephanos = Cyclosorus
Hemigramma = Tectaria Melpomene (37) Platycerium (37) Sphenomeris (23)
Hemionitis (26) Meniscium = Cyclosorus Platygyria = Lepisorus Steenisioblechnum (30)
Hemitelia = Cyathea Menisorus = Cyclosorus Platyloma (26) Stegnogramma = Cyclosorus
Heterogonium (34) Merinthosorus = Aglaomorpha Platyzoma (26) Steiropteris = Cyclosorus
Hippochaete = Equisetum Meryngium = Hymenophyllum Plecosorus = Polystichum Stenochlaena (30)
Histiopteris (25) Mesophlebion = Cyclosorus Pleocnemia (34) Stenolepia (32)
Holcochlaena (26) Metathelypteris = Thelypteris Pleopeltis (37) Sticherus (7)
Holodictyum = Asplenium Metaxya (22) Plesioneuron = Cyclosorus Stigmatopteris (32)
Holostachyum = Aglaomorpha Microgonium = Didymoglossum Pleuroderris = Tectaria Stromatopteris (7)
Homalosorus (29) Microgramma (37) Pleurosoriopsis (37) Synammia (37)
Humata = Davallia Microlepia (25) Pleurosorus = Asplenium Syngramma (26)
Hyalotricha = Campyloneurum Micropolypodium (37) Pneumatopteris = Cyclosorus Taenitis (26)
Hyalotrichopteris = Campyloneurum Microsorum (37) Podosorus (37) Tapeinidium (23)
Hymenasplenium (27) Microstaphyla = Elaphoglossum Polybotrya (32) Tectaria (34)
Hymenocystis = Woodsia Microtrichomanes = Polyphlebium (6) Teratophyllum (32)
Hymenoglossum = Hymenophyllum Hymenophyllum Polypodioides (37) Terpsichore (37)
Hymenophyllopsis (20) Mildella (26) Polypodiopteris (37) Thamnopteris = Asplenium
Hymenophyllum (6) Mohria = Anemia Polypodium (37) Thelypteris (28)
Hypodematium (32) Monachosorum (25) Polystichopsis (32) Themelium (37)
Hypoderris (34) Monogramma (26) Polystichum (32) Thylacopteris (37)
Hypolepis (25) Monomelangium = Diplazium Polytaenium (26) Thyrsopteris (15)
Idiopteris = Pteris Neocheiropteris (37) Pronephrium = Cyclosorus Thysanosoria (33)
Jamesonia (26) Nephelea = Alsophila Prosaptia (37) Tmesipteris (2)
Japanobotrychium = Botrychium Nephopteris (26) Protowoodsia (29) Todea (5)
Kontumia (37) Nephrolepis (33) Psammiosorus (34) Trachypteris (26)
Kuniwatsukia = Athyrium Neurocallis (26) Pseudocolysis = Pleopeltis Trichoneuron (32)
Lacostea = Trichomanes Neurodium (37) Pseudocyclosorus = Cyclosorus Trichipteris = Cyathea
Lacosteopsis = Vandenboschia Neuromanes = Trichomanes Pseudocystopteris (29) Trichomanes (6)
Lastrea = Thelypteris Niphidium (37) Pseudodrynaria = Aglaomorpha Trigonospora = Cyclosorus
Lastreopsis (32) Notholaena (26) Pseudophegopteris (28) Triplophyllum (34)
Lecanium = Didymoglossum Nothoperanema = Dryopteris Pseudotectaria = Tectaria Trismeria = Pityrogramma
Lecanopteris (37) Ochropteris (26) Psilotum (2) Vaginularia = Monogramma
Lellingeria (37) Odontosoria (23) Psomiocarpa (34) Vandenboschia (6)
Lemmaphyllum (37) Oenotrichia (25) Pteridium (25) Vittaria (26)
Lepisorus (37) Oenotrichia p.p (32) Pteridoblechnum (30) Weatherbya = Lemmaphyllum
Leptochilus (37) Oleandra (35) Pteridrys (34) Woodsia (29)
Leptogramma = Cyclosorus Olfersia (32) Pteris (26) Woodwardia (30)
Leptolepia (25) Onoclea (31) Pterozonium (26) Xiphopteris = Cochlidium
Leptopteris (5) Onocleopsis (31) Ptilopteris = Monachorosum Xyropteris (23)
Leptorumohra = Arachniodes Onychium (26) Pycnodoria = Pteris Zygophlebia (37)
... For a long time, pteridologists have recognized significant differences among the Matoniaceae, Dipteridaceae, and Gleicheniaceae [60], so that Matoniaceae and Dipteridaceae have been treated into the two orders, Matoniales [30] and Dipteridales [29], respectively. Soon after, Mationaceae and Dipteridaceae were merged into Gleicheniales along with Gleicheniaceae, based on monophyletic evidence [28,59,61] and some morphological traits such as root steles with 3-5 protoxylem poles [62] and antheridia with 6-12 narrow, twisted, or curved cells in walls [63], which usually are called gleichids because of common traits of forking branches. In fact, these traits are very common among early leptosporangiate ferns, such as Schizaea dichotoma (Schizaeaceae), Lygodium digitatum (Lygodiaceae), Regnellidium diphyllum (Marsileaceae), and Gonocormus minutus (Hymenophyllaceae), and therefore do not provide the classification significance for high order rank. ...
... Based on this robust evidence from phylogenomics and morphology, we propose to redefine a general ordinal rank of Matoniales from Dipteridales Doweld (2001) and Matoniales Reveal (1993) based on the International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) [68]. This order should be composed of extant Dipteridaceae (including Cheiropleuria and Dipteris) and Matoniaceae (Matonia and Phanerosorus), with a total of 15 species [20,63], which flourished with widespread global distribution throughout the Mesozoic era together with their related group of Gleicheniaceae [54,[69][70][71]. Certainly, the redefinition of order Matoniales will provide new insights regarding the diversification and prosperity of fossil genera of Matoniaceae and Dipteridaceae, such as Clathropteris, Dictyophyllum, Hausmannia, Goeppeetella, Camptopteris, Thaumatopteris, Phlebopteris, Matonidium, Weichselia, and so on [57,69,[72][73][74][75]. ...
... Only a few new orders of land organisms, such as Mantophasmatodea [6,77] and Hypopterygiales [9], have been recently published and accepted based on phylogenomic evidence. In the extant lycopods and ferns, more than 40 valid orders were reported, but only 14 of these are accepted in PPG I [20,63]. However, more careful studies may be required to verify the treatments of these synonym names in the future [78]. ...
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Phylogenetic conflicts limit our understanding of the evolution of terrestrial life under multiple whole genome duplication events, and the phylogeny of early terrestrial plants remains full of controversy. Although much incongruence has been solved with so-called robust topology based on single or lower copy genes, the evolutionary mechanisms behind phylogenetic conflicts such as polyploidization remain poorly understood. Here, through decreasing the effects of polyploidization and increasing the samples of species, which represent all four orders and eight families that comprise early leptosporangiate ferns, we have reconstructed a robust phylogenetic tree and network with 1125 1-to-1 orthologs based on both coalescent and concatenation methods. Our data consistently suggest that Matoniales, as a monophyletic lineage including Matoniaceae and Dipteridaceae, should be redefined as an ordinal rank. Furthermore, we have identified and located at least 11 whole-genome duplication events within the evolutionary history of four leptosporangiates lineages, and associated polyploidization with higher speciation rates and mass extinction events. We hypothesize that paleopolyploidization may have enabled leptosporangiate ferns to survive during mass extinction events at the end Permian period and then flourish throughout the Mesozoic era, which is supported by extensive fossil records. Our results highlight how ancient polyploidy can result in rapid species radiation, thus causing phylogenetic conflicts yet allowing plants to survive and thrive during mass extinction events.
... The first radiation occurred during the Palaeozoic (Rothwell 1999), and the second radiation can be traced back to the early Triassic (Skog 2001), where fossils assignable to modern genera Osmunda sensu lato (s.l.) were found in the Triassic of Antarctica (Phipps et al. 1998). By the Jurassic, all the basal families of living leptosporangiate ferns were produced (Skog 2001;Smith et al. 2006;Rothwell & Stockey 2008). The families represented in this radiation encompass Osmundaceae, Gleicheniaceae, Hymenophyllaceae, Dipteridaceae, Matoniaceae, Schizaeaceae s.l., and Cyatheaceae s.l. ...
... The families represented in this radiation encompass Osmundaceae, Gleicheniaceae, Hymenophyllaceae, Dipteridaceae, Matoniaceae, Schizaeaceae s.l., and Cyatheaceae s.l. (Smith et al. 2006). These families are well represented in different Middle Jurassic floras worldwide (Barale & Ouaja 2002;Wang 2002;Cleal & Rees 2003;Birkenmajer & Ociepa 2008;Mehlqvist et al. 2009;Barbacka 2011;Vaez-Javadi 2011;Kostina & Herman 2013;Scanu et al. 2015), but their distribution is far from understood in low-latitude regions such as Mexico. ...
... Our material can be distinguished from Calochlaena (Maxon) M.D. Turner & R.A. White and Dicksonia M.R. Schomb. due the lack of sori with bivalvate or cuplike indusium in a marginal position (Smith et al. 2006). It differs from fossil genera such as Coniopteris Brongniart and Eboracia Thomas due the lack of marginal sori with a cup-shaped indusium (Harris 1961;Li et al. 2020). ...
Mesozoic ferns from Mexico have been the subject of serious academic endeavours since the beginning of the 20th Century, to understand these plants at the time of their peak diversity. Most findings have been made in a set of Middle Jurassic Basins of the Mixtec Terrain in the Oaxaca State. However, fossil ferns are scarce in other assemblages, so further identification of this group has been infrequent. Here we describe six new and recently collected fossil plants from the Middle Jurassic Otlaltepec Formation, Puebla. Based on their fertile and vegetative fronds, we propose the new genus Paralophosoria Morales-Toledo, Mendoza-Ruiz & Cevallos-Ferriz, gen. nov. in the Dicksoniaceae, represented by Paralophosoria jurassica , Morales-Toledo, Mendoza-Ruiz & Cevallos-Ferriz, sp. nov., and identify the following genera: cf . Aspidistes , Sphenopteris , Spiropteris . A fern with uncertain affinities was also described. This work contributes to the understanding of fern diversity in low latitudes during the Middle Jurassic in Mexico.
... The plant macrofossil occurrences from the Los Rastros, Ischigualasto, and Los Colorados formations were organized by more inclusive taxonomic groups. This was carried out following the systematic schemes proposed by Stewart and Rothwell (1993) for gymnosperms, Smith et al. (2006), and PPG (2016) for ferns (Supplementary Table S10). For palynomorphs recorded from the same formations, an attempt was made to establish the botanical affinity of the miospores with the plant group interpreted as the most likely producer based on Ruffo Rey (2021) and references therein (Supplementary Table S11). ...
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Paleoenvironmental factors such as climate change are often hypothesized as critical for ecosystem changes over evolutionary time scales. Theoretically, these changes should be observable in the fossil record, but the robust linkage of biotic shifts to climate events remains difficult. The Late Triassic Period is an ideal interval for testing such hypotheses within terrestrial ecosystems because of a number of large-scale climate events, such as the Carnian Pluvial Episode (CPE), and a rich, well-studied fossil record. Nonetheless, at regional to global scales, few linkages have been identified during most of the Late Triassic. Here, we synthesize a multi-proxy basin-scale dataset of paleoenvironmental data, including new information from clay mineralogy and paleosol major- and trace-element geochemistry, to understand paleoclimate changes in the Ischigualasto-Villa Unión Basin of northwestern Argentina. In combination with diversity and relative abundance information from the well-sampled plant and tetrapod fossil record in the basin, we then use accepted evolutionary models of abiotic forcing of changes in ecosystems to test previously proposed hypotheses of floral and faunal variations at a basin scale. We find that although many patterns are best explained by sampling issues and taphonomy, pseudosuchian archosaur diversity and rhynchosaur relative abundance conform to predictions of paleoenvironmental forcing as the climate changed from warmer, drier conditions to more temperate humid conditions. These data demonstrate how high-resolution multi-proxy data and a well-sampled fossil record at a basin-scale can be used to test hypotheses of abiotic forcing of ecosystems in deep time, and highlights where future efforts should be directed in terms of filling data gaps and testing new hypotheses.
... The genus Taenitis Willd. ex Schkuhr was first introduced by Schkuhr (Smith et al. 2006). In 19th century, Taenitis had been classified in Adiantaceae (Holttum 1968). ...
... Molecular and fossil evidence places the origin of ferns at around 400 Ma, more than 200 Myr before the appearance of angiosperms (Kenrick & Crane, 1997;Soltis et al., 1999). The rise of angiosperms in the Cretaceous period led to darker forest understories and coincided with a decline in fern and gymnosperm species richness, but leptosporangiate ferns (Polypodiidae) (Christenhusz et al., 2011;The Pteridophyte Phylogeny Group, 2016) subsequently proliferated to become the second most species-rich group of vascular plants, and now comprise the vast majority of extant fern species (Schneider et al., 2004;Smith et al., 2006;Schuettpelz & Pryer, 2009). ...
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Ferns appear in the fossil record some 200 Myr before angiosperms. However, as angiosperm-dominated forest canopies emerged in the Cretaceous period there was an explosive diversification of modern (leptosporangiate) ferns, which thrived in low, blue-enhanced light beneath angiosperm canopies. A mechanistic explanation for this transformative event in the diversification of ferns has remained elusive. We used physiological assays, transcriptome analysis and evolutionary bioinformatics to investigate a potential connection between the evolution of enhanced stomatal sensitivity to blue light in modern ferns and the rise of angiosperm-dominated forests in the geological record. We demonstrate that members of the largest subclade of leptosporangiate ferns, Polypodiales, have significantly faster stomatal response to blue light than more ancient fern lineages and a representative angiosperm. We link this higher sensitivity to levels of differentially expressed genes in blue-light signaling, particularly in the cryptochrome (CRY) signaling pathway. Moreover, CRYs of the Polypodiales examined show gene duplication events between 212.9–196.9 and 164.4–151.8 Ma, when angiosperms were emerging, which are lacking in other major clades of extant land plants. These findings suggest that evolution of stomatal blue-light sensitivity helped modern ferns exploit the shady habitat beneath angiosperm forest canopies, fueling their Cretaceous hyperdiversification.
The genus Lepisorus is often associated with complex lineages in Polypodiaceae, which leads to difficulty in understanding taxonomic relationships among the species. The taxa is mostly epiphytic with some distinct features like rhizomes covered with clathrate scales, simple fronds, sori in a single row, intermixed with peltate paraphyses. In this study, we precisely focus on the Lepisorus species and seek a broader understanding of the taxonomic relationships that prevail in this genus. The morpho-anatomical traits of rhizomes, stipe, lamina, and sporangia were explored in nine species with the aid of light microscopy and scanning electron microscopy techniques for gathering the qualitative and quantitative data sets. These character traits were resolved into groups based on degree of similarity and principal component analysis to comprehend the covariance among the variables. The data were then employed to prepare an artificial dichotomous key and hierarchical cluster analysis was established that revealed five related clades with eight members whereas one member emerged as outgroup. The results correspond with other recent phylogenetic studies of the concerned genus hence confirming immense reliability and thrust of light microscopy, scanning electron microscopy, and morphology-based studies that are being less utilized in fern taxonomy.
Adder's tongue ferns or Ophioglossaceae are best known among evolutionary biologists and botanists for their highest chromosome count of any known organisms, the presence of sporophores, and simple morphology. Previous studies recovered and strongly supported the monophyly of the family and the two multi-generic subfamilies, Botrychioideae and Ophioglossoideae, but the relationships among these and two other subfamilies (Helminthostachyoideae and Mankyuoideae) are not well resolved preventing us from understanding the character evolution. The monophyly of and the relationships in the species-rich genus, Ophioglossum, have not well been understood. In this study, new phylogenetic trees are reconstructed based on four datasets: Sanger sequences of eight plastid markers of 184 accessions, 22 plastomes (12 are new), 29 morphological characters, and combined Sanger and morphological data. Our major results include: (1) the relationships among the four subfamilies are well resolved and strongly supported in Bayesian and parsimony analyses based on plastomes: Mankyua is sister to the rest, followed by Ophioglossoideae which are sister to Helminthostachys + Botrychioideae; (2) Sanger data, plastomes, and combined Sanger and morphological data recovered and strongly supported the monophyly of Ophioglossum in its current circumscription (sensu lato; s.l.) in Bayesian and/or parsimony analyses; (3) within Ophioglossum s.l., four deeply diverged clades are identified and the relationships among the four clades are well resolved; (4) evolution of 34 morphological characters is analyzed in the context of the new phylogeny, among which shape of rhizomes, germination time of spores, shape of early gametophytes, and a number of other characters are found to contain interesting phylogenetic signal; and (5) based on the new phylogeny and character evolution, we propose a new classification of Ophioglossaceae in which the currently circumscribed Ophioglossum is divided into four genera including three new ones: Goswamia, Haukia, and Whittieria considering their molecular, morphological, ecological, and biogeographical distinctiveness.
Background Matteuccia struthiopteris (L.) Todaro is a popular edible wild fern found in the subtropical Himalayas. It is widely used in folk medicine to treat several ailments and known for its antioxidant, anti-inflammatory, antibacterial, antiviral and antidiabetic activities. Due to its multidimensional nutritional, pharmacological and therapeutic effects, it is well recognized in the Ayurvedic Pharmacopoeia. Main body of abstract The present review aims to provide updated information on Matteuccia struthiopteris botany, phytochemistry, pharmacological effects and toxicity methods, in addition to highlight potential for future exploration. Particular emphasis is also given to its antioxidant potential in health promotion. In-depth literature was probed by examining numerous sources via online databases (research and reviews), texts, Web sites and thesis. Plant biotechnology approaches such as tissue culture and micropropagation are also discussed. Short conclusion Matteuccia struthiopteris is found useful in the treatment of different diseases such as microbial infection, viral infection and diabetes and in weight reduction and also effective as antioxidant and free radical scavenger. Nevertheless, advance studies are required to offer the mechanistic role of crude extracts and its bio-actives and even to discover the structure–function relationship of active principles.
While continuing to be developed as a fern model, Ceratopteris richardii has demonstrated its value in elucidating evolutionary developmental questions in land plants, in addition to other aspects of plant biology. Plants are unique in that they have the ability to produce new organs throughout their life due to persistent proliferation of tissue-specific stem cell populations. The family of WUSCHEL-related homeobox (WOX) genes are important transcription factors for stem cell maintenance. WOX genes are found in all land plants and some extant algae. This chapter concentrates on the current understanding of the evolution of the WOX gene family in controlling cell proliferation of various meristem domains in distantly related land plant models, Physcomitrella patens, C. richardii, and Arabidopsis thaliana.
First record is made of a dioecious condition in the heterosporous, terrestrial fern Platyzoma microphyllum R. Br. from northern Australia. The spores, gametophytes and young sporophyte are described and the role of heterospory and dioecism in these plants is discussed. The genus is newly classified as a subfamily Platyzomatoideae in the Polypodiaceae.
Two subfamilies, four genera, with 35 species of the fern family Woodsiaceae (Diels) Herter are recognised and enumerated with synonyms and distribution from temperate and tropical mountain areas of Eurasia. Three new combinations are made: subgen. Acrolysis (Nakai) Shmakov, comb. et stat. nov., subgen. Eriosorus (Ching) Shmakov, comb. et stat. nov., Cheilanthopsis kangdingensis (H. S. Kung, L.B. Zhang and X.S. Guo) Shmakov, comb. nov. One new series is described: Glabellae Shmakov, ser. nov. Key to the genera and species is also provided.
The new genus Kontumia (Polypodiaceae?) and its type species Kontumia heterophylla from Vietnam are described and illustrated. Kontumia is distinguished from Gymnogrammitis by its scandent habit, dimorphic leaves, clathrate transparent scales on the rhizome and petiole bases, and aspects of leaf morphology. Also, a new species of Cyrtomium from Vietnam is described and named as C. elongatum (Dryopteridaceae). Cyrtomium elongatum is similar to C. wulingense S. F. Wu, but differs by its longer leaves (to 90 cm) and more numerous pinnae (more than 60 pairs) with cordate rather than oblique bases, abaxially with sparse small brown scales, and with brown punctations within the areoles.