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A Classification for Extant Ferns

August 2006
Taxon 55(3):705-731

August 2006
55(3):705-731

DOI:10.2307/25065646

Authors:

Kathleen Pryer

Duke University

Eric Schuettpelz

Smithsonian Institution

Petra Korall

Uppsala University

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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.

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INTRODUCTION

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-

705

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

≥70.

Euphyllophytes

MonilophytesSpermatophytesLycophytes

TAXONOMY

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. arsmith@berkeley.edu (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.

KEYWORDS:

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

706

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

future.

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 (http://www.life.umd.edu/emeritus/reveal/

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).

CLASSIFICATION OF EXTANT

FERNS

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

707

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.

Psilotaceae

Ophioglossaceae Psilotopsida

Psilotales

Ophioglossales

EquisetopsidaEquisetalesEquisetaceae

MarattiopsidaMarattialesMarattiaceae

Polypodio-

psida

OsmundalesOsmundaceae

Gleicheniales

Gleicheniaceae

Dipteridaceae

Matoniaceae

Hymenophyllales

Hymenophyllaceae

Schizaeales

Schizaeaceae

Lygodiaceae

Anemiaceae

Salviniales

Salviniaceae

Marsileaceae

Cyatheales

Plagiogyriaceae

Loxomataceae

Thyrsopteridaceae

Cibotiaceae

Cyatheaceae

Polypodiales

Lindsaeaceae

Saccolomataceae

Dennstaedtiaceae

Pteridaceae

Aspleniaceae

Thelypteridaceae

Woodsiaceae

Onocleaceae

Blechnaceae

Dryopteridaceae

Davalliaceae

Polypodiaceae

Leptosporangiates

Core leptosporangiates

Ferns

Polypods

Eupolypods

Eupolypods II

Eupolypods I

Metaxyaceae

Dicksoniaceae

Culcitaceae

Oleandraceae

Tectariaceae

Lomariopsidaceae

708

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.

I. CLASS PSILOTOPSIDA

A. ORDER OPHIOGLOSSALES.

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).

B. ORDER PSILOTALES.

2. Family Psilotaceae. — Whisk ferns; incl.

Tmesipteridaceae.

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.

II. CLASS EQUISETOPSIDA [= SPHE-

NOPSIDA].

C. ORDER EQUISETALES.

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.

III. CLASS MARATTIOPSIDA

D. ORDER MARATTIALES. — Including

Christenseniales.

4. Family Marattiaceae. — Marattioids; incl.

Angiopteridaceae, Christenseniaceae, Danaeaceae,

Kaulfussiaceae.

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

sense.

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

709

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).

IV. CLASS POLYPODIOPSIDA [= FI-

LICOPSIDA]

E. ORDER OSMUNDALES.

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.

F. ORDER HYMENOPHYLLALES.

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.

G. ORDER GLEICHENIALES. — Including

Dipteridales, Matoniales, Stromatopteridales.

Mo-

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-

pteridaceae.

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-

pleuriaceae.

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

710

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.

Two

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).

H. ORDER SCHIZAEALES. —

Monophyletic

(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-

ceae.

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,

103.

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

711

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

(Azolla).

J. ORDER CYATHEALES. —

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

712

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.

K. ORDER POLYPODIALES. —

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

713

gametophytes green, usually cordate (sometimes ribbon-

shaped in some epiphytes), surficial.

23. Family Lindsaeaceae. —

Lindsaeoids; incl.

Cystodiaceae, Lonchitidaceae. Ca. eight genera: Cystodi-

um,Lindsaea,Lonchitis,Odontosoria,Ormoloma

,Sphe-

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-

ers.

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. —

Dennstaedt-

ioids; incl. Hypolepidaceae, Monachosoraceae, Pteridia-

ceae. Ca. 11 genera: Blotiella, Coptodipteris, Denn-

staedtia (incl. Costaricia

), Histiopteris, Hypolepis,

Leptolepia, Microlepia, Monachosorum, Oenotrichia

s.s.

, 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-

sora

, 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

714

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. —

Asplenioids,

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. —

Thelypteroids

or thelypteridoids; incl. “Sphaerostephanaceae”. Circa

Smith & al. • Fern classification55 (3) • August 2006: 705–731

715

5–30 genera, depending on taxonomic viewpoint: com-

monly accepted segregates are Cyclosorus (incl. Ampel-

opteris

, 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

morphologically.

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-

tis

; 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-

pteris).

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

716

& 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. —

Dryopteroids

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-

tsia

(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

Dryopteridaceae.

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 (http://www.nybg.org/bsci/res/moran/ela

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

717

Dryopteridaceae is certainly paraphyletic.

33. Family Lomariopsidaceae. —

Lomariop-

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:

Aenigmopteris

, Arthropteris, Heterogonium, Hypoder-

ris

, Pleocnemia, Psammiosorus, Psomiocarpa

, Pteri-

drys, Tectaria s.l. (incl. Amphiblestra

,Camptodium

Chlamydogramme

, Cionidium, Ctenitopsis, Dictyo-

xiphium, Fadyenia, Hemigramma, Pleuroderris

, Pseu-

dotectaria

, 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

718

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,

Gymnogrammitis,Kontumia

(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).

ACKNOWLEDGEMENTS

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

719

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.

LITERATURE CITED

Angiosperm Phylogeny Group [APG II]. 2003. An update of

the Angiosperm Phylogeny Group classification for the

orders and families of flowering plants: APG II. Bot. J.

Linn. Soc. 141: 399–436.

Barcelona, J. F. 2000. Systematics of the Fern Genus Odonto-

soria Sensu Lato (Lindsaeaceae). Ph.D. thesis, Miami

Univ., Oxford, Ohio.

Bateman, R. M. 1991. Paleobiological and phylogenetic im-

plications of anatomically-preserved Archeocalamites

from the Dinantian of Oxroad Bay and Loch Humphrey

Burn, southern Scotland. Palaeontographica B223: 1–59.

Beckert, S., Muhle, H., Pruchner, D. & Knoop, V. 1999. Phy-

logenies from mitochondrial DNA in cryptogams. Pp. 73

in: Stuessy, T. F. & Greimler, J. (eds.), XVII International

Botanical Congress, Vienna, Abstracts book. Organizing

Committee, Vienna.

Bower, F. O. 1926. The Ferns (Filicales). Vol. 2. The Euspo-

rangiate and Other Relatively Primitive Ferns. Cambridge

Univ. Press, London.

Brummitt, R. K. 1996. In defence of paraphyletic taxa. Pp.

371–384 in: Maesen, L. J. G., van der Burgt, X. M. & van

Medenbach var Rooy, J. M. (eds.), The Biodiversity of

African Plants. Proceedings XIVth AETFAT Congress

22–27 Aug. 1994. Kluwer Academic Publishers, Nether-

lands.

Brummitt, R. K. 1997. Taxonomy versus cladonomy, a funda-

mental controversy in biological systematics. Taxon 46:

723–734.

Ching, R. C. 1940. On natural classification of the family

“Polypodiaceae”. Sunyatsenia 5: 201–268.

Ching, R. C. 1978. The Chinese fern families and genera: sys-

tematic arrangement and historical origin. Acta Phytotax.

Sin. 16 (3): 1–19; 16 (4): 16–37.

Christensen, C. 1938. Filicinae. Pp. 522–550 in: Verdoorn, F.

(ed.), Manual of Pteridology. Martinus Nijhoff, The

Hague.

Collinson, M. E. 1996. “What use are fossil ferns?”—20 years

on: with a review of the fossil history of extant pterido-

phyte families and genera. Pp. 349–394 in: Camus, J. M.,

Gibby, M. & Johns, R. J. (eds.), Pteridology in Perspec-

tive. Royal Botanic Gardens, Kew.

Conant, D. S., Raubeson, L. A., Attwood, D. K., Perera, S.,

Zimmer, E. A., Sweere, J. A. & Stein, D. B. 1996. Phy-

logenetic and evolutionary implications of combined anal-

ysis of DNA and morphology in the Cyatheaceae. Pp.

231–248 in: Camus, J. M., Gibby, M. & Johns, R. J. (eds.),

Pteridology in Perspective. Royal Botanic Gardens, Kew.

Conant, D. S., Raubeson, L. A., Attwood, D. K. & Stein, D.

B. 1996 [“1995”]. The relationships of Papuasian Cyathe-

aceae to New World tree ferns. Amer. Fern J. 85: 328–340.

Conant, D. S. & Stein, D. B. 2001. Phylogenetic and geo-

graphic relationships of the tree ferns (Cyatheaceae) on

Mount Kinabalu. Sabah Parks Nat. J. 4: 25–43.

Copeland, E. B. 1947. Genera Filicum. Chronica Botanica,

Waltham, Mass.

Crabbe, J. A., Jermy, A. C. & Mickel, J. T. 1975. A new ge-

neric sequence for the pteridophyte herbarium. Fern Gaz.

11: 141–162.

Crane, E. H. 1997. A revised circumscription of the genera of

the fern family Vittariaceae. Syst. Bot. 22: 509–517.

Crane, E. H., Farrar, D. R. & Wendel, J. F. 1996 [“1995”].

Phylogeny of the Vittariaceae: convergent simplification

leads to a polyphyletic Vittaria. Amer. Fern J. 85:

283–305.

Cranfill, R. B. 2001. Phylogenetic Studies in the Polypodiales

(Pteridophyta) with an Emphasis on the Family

Blechnaceae. Ph.D. thesis, Univ. California, Berkeley.

Cranfill, R. B. & Kato, M. 2003. Phylogenetics, biogeogra-

phy and classification of the woodwardioid ferns

(Blechnaceae). Pp. 25–48 in: Chandra, S. & Srivastava, M.

(eds.), Pteridology in the New Millennium. Kluwer

Academic Publishers, Dordrecht.

Crisp, M. D., Gilmore, S. R. & Weston, P. H. 1999. Phylog-

enetic relationships of two anomalous species of Pulte-

naea (Fabaceae: Mirbelieae), and description of a new

genus. Taxon 48: 701–714.

Croft, J. R. 1986. The stipe and rachis vasculature of the dick-

sonioid fern, Cystodium sorbifolium (Cystodiaceae). Kew

Bull. 41: 789–803.

Davidse, G., Sousa S., M. & Knapp, S. (general eds.). 1995.

Flora Mesoamericana. Vol. 1. Psilotaceae a Salviniaceae.

(Moran, R. C. & Riba, R., vol. eds.). Universidad Nacional

Autónoma de México, México, D.F.

Davies, K. L. 1991. A brief comparative survey of aerophore

structure within the Filicopsida. Bot. J. Linn. Soc. 197:

115–137.

Des Marais, D. L., Smith, A. R., Britton, D. M. & Pryer, K.

M. 2003. Phylogenetic relationships and evolution of ex-

tant horsetails, Equisetum, based on chloroplast DNA se-

quence data (rbcL and trnL-F). Int. J. Pl. Sci. 164:

737–751.

Dickason, F. G. 1946. The ferns of Burma. Ohio J. Sci. 46:

109–141.

Diggs Jr., G. M. & Lipscomb, B. L. 2002. What is the writer

of a flora to do? Evolutionary taxonomy or phylogenetic

systematics? Sida 20: 647–674.

Dubuisson, J.-Y. 1996. Evolutionary relationships within the

genus Trichomanes sensu lato (Hymenophyllaceae) based

on anatomical and morphological characters and a com-

parison with rbcL nucleotide sequences; preliminary

results. Pp. 285–287 in: Camus, J. M., Gibby, M. & Johns,

R. J. (eds.), Pteridology in Perspective. Royal Botanic

Gardens, Kew.

Dubuisson, J.-Y. 1997. rbcL sequences: a promising tool for

the molecular systematics of the fern genus Trichomanes

(Hymenophyllaceae)? Molec. Phylog. Evol. 8: 128–138.

Dubuisson, J.-Y., Hennequin, S., Douzery, E. J. P., Cranfill,

R. B., Smith, A. R. & Pryer, K. M. 2003. rbcL phyloge-

ny of the fern genus Trichomanes (Hymenophyllaceae),

with special reference to neotropical taxa. Int. J. Pl. Sci.

164: 753–761.

Smith & al. • Fern classification 55 (3) • August 2006: 705–731

720

Ebihara, A., Dubuisson, J.-Y., Iwatsuki, K., Hennequin, S.

& Ito, M. In press. A taxonomic revision of Hymenophyl-

laceae. Blumea 51.

Ebihara, A., Hennequin, S., Iwatsuki, K., Bostock, P. D.,

Matsumoto, S., Jaman, R., Dubuisson, J.-Y. & Ito, M.

2004. Polyphyletic origin of Microtrichomanes (Prantl)

Copel. (Hymenophyllaceae), with a revision of the

species. Taxon 53: 935–948.

Ebihara, A., Iwatsuki, K., Kurita, S. & Ito, M. 2002.

Systematic position of Hymenophyllum rolandi-principis

Rosenst. or a monotypic genus Rosenstockia Copel.

(Hymenophyllaceae) endemic to New Caledonia. Acta

Phytotax. Geobot. 53: 35–49.

Gastony, G. J. 1982. Spore morphology of the Dicksoniaceae.

2. The genus Cibotium. Canad. J. Bot. 60: 955–972.

Gastony, G. J. & Johnson, W. P. 2001. Phylogenetic place-

ments of Loxoscaphe thecifera (Aspleniaceae) and

Actiniopteris radiata (Pteridaceae) based on analysis of

rbcL nucleotide sequences. Amer. Fern J. 91: 197–213.

Gastony, G. J. & Rollo, D. R. 1995 [1996]. Phylogeny and

generic circumscriptions of cheilanthoid ferns

(Pteridaceae: Cheilanthoideae) inferred from rbcL

nucleotide sequences. Amer. Fern J. 85: 341–360.

Gastony, G. J. & Rollo, D. R. 1998. Cheilanthoid ferns

(Pteridaceae: Cheilanthoideae) in the southwestern United

States and adjacent Mexico—a molecular phylogenetic

reassessment of generic lines. Aliso 17: 131–144.

Gastony, G. J. & Ungerer, M. C. 1997. Molecular systematics

and a revised taxonomy of the onocleoid ferns

(Dryopteridaceae: Onocleeae). Amer. J. Bot. 84: 840–849.

Gastony, G. J. & Yatskievych, G. 2001. Morphological

reassessment of molecular phylogenetic inferences in the

xerically adapted cheilanthoid ferns (Pteridaceae: Cheil-

anthoideae). Pp. 82–83 in: Botany 2001, Albuquerque,

New Mexico. New Mexico.

Geiger, J. M. O. & Ranker, T. A. 2005. Molecular phyloge-

netics and historical biogeography of Hawaiian Dryopteris

(Dryopteridaceae). Molec. Phylog. Evol. 34: 392–407.

Grant, V. 2003. Incongruence between cladistic and taxonom-

ic systems. Amer. J. Bot. 90: 1263–1270.

Greuter, W., McNeill, J., Barrie, F. R., Burdet, H. M., De-

moulin, V., Filgueiras, T. S., Nicolson, D. H., Silva, P.

C., Skog, J. E., Trehane, P., Turland, N. J. & Hawks-

worth, D. L. (eds.). 2000. International Code of Botanical

Nomenclature (Saint Louis Code) Adopted by the Six-

teenth International Botanical Congress St. Louis, Mis-

souri, July–August 1999. Koeltz Scientific Books, König-

stein. [Regnum Veg. 138.]

Guillon, J. M. 2004. Phylogeny of horsetails (Equisetum)

based on the chloroplast rps4 gene and adjacent noncod-

ing sequences. Syst. Bot. 29: 251–259.

Hasebe, M., Omori, T., Nakazawa, M., Sano, T., Kato, M. &

Iwatsuki, K. 1994. rbcL gene sequences provide evidence

for the evolutionary lineages of leptosporangiate ferns.

Proc. Natl. Acad. Sci. U.S.A. 91: 5730–5734.

Hasebe, M., Wolf, P. G., Pryer, K. M., Ueda, K., Ito, M.,

Sano, R., Gastony, G. J., Yokoyama, J., Manhart, J. R.,

Murakami, N., Crane, E. H., Haufler, C. H. & Hauk,

W. D. 1995 [1996]. Fern phylogeny based on rbcL

nucleotide sequences. Amer. Fern J. 85: 134–181.

Haufler, C. H., Grammer, W. A., Hennipman, E., Ranker, T.

A., Smith, A. R. & Schneider, H. 2003. Systematics of

the ant-fern genus Lecanopteris (Polypodiaceae): testing

phylogenetic hypotheses with DNA sequences. Syst. Bot.

28: 217–227.

Hauk, W. D. 1995 [1996]. A molecular assessment of relation-

ships among cryptic species of Botrychium subgenus Bo-

trychium (Ophioglossaceae). Amer. Fern J. 85: 375–394.

Hauk, W. D., Parks, C. R. & Chase, M. W. 2003. Phyloge-

netic studies of Ophioglossaceae: evidence from rbcL and

trnL-F plastid DNA sequences and morphology. Molec.

Phylog. Evol. 28: 131–51.

Hennequin, S., Ebihara, A., Ito, M., Iwatsuki, K. &

Dubuisson, J.-Y. 2003. Molecular systematics of the fern

genus Hymenophyllum s.l. (Hymenophyllaceae) based on

chloroplastic coding and noncoding regions. Molec.

Phylog. Evol. 27: 283–301.

Hennipman, E. 1996. Scientific consensus classification of

Pteridophyta. Pp. 191–202 in: Camus, J. M., Gibby, M. &

Johns, R. J. (eds.), Pteridology in Perspective. Royal

Botanic Gardens, Kew.

Hill, C. R. & Camus, J. M. 1986. Evolutionary cladistics of

marattialean ferns. Bull. Brit. Mus. (Nat. Hist.), Bot. 14:

219–300.

Holttum, R. E. 1947. A revised classification of leptosporan-

giate ferns. J. Linn. Soc., Bot. 53: 123–158.

Holttum, R. E. 1949. The classification of ferns. Biol. Rev. 24:

267–296.

Holttum, R. E. 1971. Studies in the family Thelypteridaceae

III. A new system of genera in the Old World. Blumea 19:

17–52.

Holttum, R. E. 1973. Posing the problems. Pp. 1–10 in: Jermy,

A. C., Crabbe, J. A. & Thomas, B. A. (eds.), The

Phylogeny and Classification of the Ferns. Academic

Press, London. [Bot. J. Linn. Soc. 67(suppl. 1)].

Holttum, R. E. 1986. Studies in the genera allied to Tectaria

Cav., V. Triplophyllum, a new genus of Africa and

America. Kew Bull. 41: 237–260.

Holttum, R. E. & Edwards, P. 1983. The tree ferns of Mt. Ro-

raima and neighboring areas of the Guayana Highlands

with comments on the family Cyatheaceae. Kew Bull. 38:

155–188.

Hoogland, R. D. & Reveal, J. L. 2005. Index nominum famil-

iarum plantarum vascularium. Bot. Rev. 71: 1–291.

Janssen, T. & Schneider, H. 2005. Exploring the evolution of

humus collecting leaves in drynarioid ferns (Polypodiace-

ae, Polypodiidae) based on phylogenetic evidence. Pl.

Syst. Evol. 252: 175–197.

Judd, W., Campbell, C. S., Kellogg, E. A., Stevens, P. F. &

Donoghue, M. J. 2002. Plant Systematics: A Phylogenetic

Approach. Ed. 2. Sinauer Associates, Sunderland,

Massachusetts.

Kato, M. 1983. The classification of major groups of pterido-

phytes. J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 13: 263–283.

Kato, M. 1987. A phylogenetic classification of Ophioglossa-

ceae. Gard. Bull. Singapore 40: 1–14.

Kato, M. & Setoguchi, H. 1998. An rbcL-based phylogeny

and heteroblastic leaf morphology of Matoniaceae. Syst.

Bot. 23: 391–400.

Kato, M., Yatabe, Y., Sahashi, N. & Murakami, N. 2001.

Taxonomic studies of Cheiropleuria (Dipteridaceae).

Blumea 46: 513–525.

Kenrick, P. & Crane, P. R. 1997. The Origin and Early

Diversification of Land Plants. Smithsonian Institution

Smith & al. • Fern classification55 (3) • August 2006: 705–731

721

Press, Washington, D.C.

Korall, P., Conant, D. S., Schneider, H., Ueda, K., Nishida,

H. & Pryer, K. M. In press. On the phylogenetic position

of Cystodium: It’s not a tree fern—it’s a polypod! Amer.

Fern J. 96(2).

Korall, P., Pryer, K. M., Metzgar, J. S., Schneider, H. & Co-

nant, D. S. 2006. Tree ferns: monophyletic groups and

their relationships as revealed by four protein-coding plas-

tid loci. Molec. Phylog. Evol. 39: 830–845.

Kranz, H. D. & Huss, V. A. R. 1996. Molecular evolution of

pteridophytes and their relationship to seed plants: evi-

dence from complete 18S rRNA gene sequences. Pl. Syst.

Evol. 202: 1–11.

Kreier, H.-P. & Schneider, H. 2006. Phylogeny and biogeog-

raphy of the staghorn fern genus Platycerium

(Polypodiaceae, Polypodiidae). Amer. J. Bot. 93: 217–225.

Kreier, H.-P. & Schneider, H. In press. Reinstatement of

Loxogramme dictyopteris for a New Zealand endemic fern

known as Anarthropteris lanceolata based on phylogenet-

ic evidence. Austral. Syst. Bot.

Kubitzki, K. (ed.). 1990. The Families and Genera of Vascular

Plants. Vol. 1. Pteridophytes and Gymnosperms. (Vol. eds.

Kramer, K. U. & Green, P. S.). Springer-Verlag, Berlin.

Lantz, T. C., Rothwell, G. W. & Stockey, R. A. 1999. Conan-

tiopteris schuchmanii, gen. et sp. nov., and the role of fos-

sils in resolving the phylogeny of Cyatheaceae s.l. J. Pl.

Res. 112: 361–381.

Lehnert, M., Mönnich, M., Pleines, T., Schmidt-Lebuhn, A.

& Kessler, M. 2001. The relictual fern genus Loxsomop-

sis. Amer. Fern J. 91: 13–24.

Lidén, M., Oxelman, B., Backlund, A., Andersson, L.,

Bremer, B., Eriksson, R., Moberg, R., Nordal, I.,

Persson, K., Thulin, M. & Zimmer, B. 1997. Charlie is

our darling. Taxon 46: 735–738.

Little, D. P. & Barrington, D. S. 2003. Major evolutionary

events in the origin and diversification of the fern genus

Polystichum (Dryopteridaceae). Amer. J. Bot. 90: 508–

514.

Lu, J.-M., Li, D.-Z, Gao, L.-M. & Cheng, X. 2005. Paraphyly

of Cyrtomium (Dryopteridaceae): evidence from rbcL and

trnL-F sequence data. J. Pl. Res. 118: 129–135.

Manhart, J. R. 1994. Phylogenetic analysis of green plant

rbcL sequences. Molec. Phylog. Evol. 3: 114–127.

Manhart, J. R. 1996 [“1995”]. Chloroplast 16S rDNA

sequences and phylogenetic relationships of fern allies and

ferns. Amer. Fern J. 85: 182–192.

Mehra, P. N. 1961. Cytological evolution in ferns with partic-

ular reference to Himalayan forms. Proc. 48th Indian Sci.

Congr. Assoc. 2: 1–24.

Mickel, J. T. 1974. Phyletic lines in modern ferns. Ann.

Missouri Bot. Gard. 61: 474–482.

Mickel, J. T. & Atehortúa, L. G. 1980. Subdivision of the

genus Elaphoglossum. Amer. Fern J. 70: 47–68.

Moore, G. 1998. A comparison of traditional and phylogenetic

nomenclature. Taxon 47: 561–579.

Murakami, N. 1995. Systematics and evolutionary biology of

the fern genus Hymenasplenium (Aspleniaceae). J. Pl.

Res. 108: 257–268.

Murakami, N., Nogami, S., Watanabe, M. & Iwatsuki, K.

1999. Phylogeny of Aspleniaceae inferred from rbcL

nucleotide sequences. Amer. Fern J. 89: 232–243.

Murakami, N. & Schaal, B. A. 1994. Chloroplast DNA varia-

tion and the phylogeny of Asplenium sect. Hymenaspleni-

um (Aspleniaceae) in the New World tropics. J. Pl. Res.

107: 245–251.

Murdock, A. 2005. Molecular Evolutionand Phylogeny of Ma-

rattioid Ferns, an Ancient Lineage of Land Plants. http://

www.2005.botanyconference.org/engine/search/index.php

Nagalingum, N. S., Schneider, H. & Pryer, K. M. In press.

Comparative morphology of reproductive structures in

heterosporous water ferns and a re-evaluation of the

sporocarp. Int. J. Plant Sci.

Nakahira, Y. 2000. A Molecular Phylogenetic Analysis of the

Family Blechnaceae, Using the Chloroplast Gene rbcL.

M.S. thesis, Graduate School of Science, Univ. Tokyo,

Tokyo.

Nakazato, T. & Gastony, G. J. 2001. Molecular phylogenetics

of Anogramma species and related genera (Pteridaceae:

Taenitidoideae). P. 86 in: Botany 2001, Albuquerque, New

Mexico. New Mexico.

Nayar, B. K. 1970. A phylogenetic classification of the homo-

sporous ferns. Taxon 19: 229–236.

Olmstead, R. G., DePamphilis, C. W., Wolfe, A. D., Young,

N. D., Elisons, W. J. & Reeves, P. A. 2001. Disintegration

of the Scrophulariaceae. Amer. J. Bot. 88: 348–361.

Pahnke, J., Goremykin, V., Bobrova, V., Troitsky, A.,

Antonov, A. & Martin, W. 1996. Utility of rDNA internal

transcribed spacer sequences from the inverted repeat of

chloroplast DNA in pteridophyte molecular phylogenetics.

Pp. 217–230 in: Camus, J. M., Gibby, M. & Johns, R. J.

(eds.), Pteridology in Perspective. Royal Botanic Gardens,

Kew.

Palmer, D. D. 1994. The Hawaiian species of Cibotium. Amer.

Fern J. 84: 73–85.

Perrie, L. R. & Brownsey, P. J. 2005. Insights into the bio-

geography and polyploid evolution of New Zealand

Asplenium from chloroplast DNA sequence data. Amer.

Fern J. 95: 1–21.

Pichi Sermolli, R. E. G. 1958. The higher taxa of the

Pteridophyta and their classification. Pp. 70–90 in:

Hedberg, O. (ed.), Systematics of today. Uppsala Univ.

Årsskrift 1958(6): 70–90.

Pichi Sermolli, R. E. G. 1959. Pteridophyta. Pp. 421–493 in:

Turrill, W. B. (ed.), Vistas in Botany. Pergamon Press Ltd.,

London.

Pichi Sermolli, R. E. G. 1965. Adumbratio Florae Aethiopicae,

11. Oleandraceae. Webbia 20: 745–769.

Pichi Sermolli, R. E. G. 1970. A provisional catalogue of the

family names of living pteridophytes. Webbia 25: 219–297.

Pichi Sermolli, R. E. G. 1973. Historical review of the higher

classification of the Filicopsida. Pp. 11–40 in: Jermy, A.

C., Crabbe, J. A. & Thomas, B. A. (eds.), The Phylogeny

and Classification of the Ferns. Academic Press, London.

[Bot. J. Linn. Soc. 67(suppl. 1)].

Pichi Sermolli, R. E. G. 1977. Tentamen pteridophytorum

genera in taxonomicum ordinem redigendi. Webbia 31:

313–512.

Pichi Sermolli, R. E. G. 1981. Report of the Subcommittee for

family names of Pteridophyta. Taxon 30: 163–168.

Pichi Sermolli, R. E. G. 1982. A further contribution to the

nomenclature of the families of Pteridophyta. Webbia 35:

223–237.

Pichi Sermolli, R. E. G. 1986. Report of the Subcommittee for

family names of Pteridophyta. Taxon 35: 686–691.

Smith & al. • Fern classification 55 (3) • August 2006: 705–731

722

Pichi Sermolli, R. E. G. 1993. New studies on some family

names of Pteridophyta. Webbia 47: 121–143.

Pinter, I., Bakker, F., Barrett, J., Cox, C., Gibby, M., Hen-

derson, S., Morgan-Richards, M., Rumsey, F., Russell,

S., Trewick, S., Schneider, H. & Vogel, J. 2002. Phylog-

enetic and biosystematic relationships in four highly dis-

junct polyploid complexes in the subgenera Ceterach and

Phyllitis in Asplenium (Aspleniaceae). Org. Diver. Evol. 2:

299–311.

Pryer, K. M. 1999. Phylogeny of marsileaceous ferns and rela-

tionships of the fossil Hydropteris pinnata reconsidered.

Int. J. Pl. Sci. 160: 931–954.

Pryer, K. M., Schneider, H. & Magallón, S. 2004a. The radi-

ation of vascular plants. Pp. 138–153 in: Cracraft, J. &

Donoghue, M. J. (eds.), Assembling the Tree of Life.

Oxford Univ. Press, New York.

Pryer, K. M., Schneider, H., Smith, A. R., Cranfill, R., Wolf,

P. G., Hunt, J. S. & Sipes, S. D. 2001a. Horsetails and

ferns are a monophyletic group and the closest living rela-

tives to seed plants. Nature 409: 618–622.

Pryer, K. M., Schuettpelz, E., Wolf, P. G., Schneider, H.,

Smith, A. R. & Cranfill, R. 2004b. Phylogeny and evolu-

tion of ferns (monilophytes) with a focus on the early lep-

tosporangiate divergences. Amer. J. Bot. 91: 1582–1598.

Pryer, K. M., Smith, A. R., Hunt, J. S. & Dubuisson, J.-Y.

2001b. rbcL data reveal two monophyletic groups of filmy

ferns (Filicopsida: Hymenophyllaceae). Amer. J. Bot. 88:

1118–1130.

Pryer, K. M., Smith, A. R. & Skog, J. E. 1996 [“1995”].

Phylogenetic relationships of extant ferns based on evi-

dence from morphology and rbcL sequences. Amer. Fern

J. 85: 205–282.

Ranker, T. A., Geiger, J. M. O., Kennedy, S. C., Smith, A. R.,

Haufler, C. H. & Parris, B. S. 2003. Molecular phyloge-

netics and evolution of the endemic Hawaiian genus

Adenophorus (Grammitidaceae). Molec. Phylog. Evol. 26:

337–347.

Ranker, T. A., Smith, A. R., Parris, B. S., Geiger, J. M. O.,

Haufler, C. H., Straub, S. C. K. & Schneider, H. 2004.

Phylogeny and evolution of grammitid ferns (Gram-

mitidaceae): a case of rampant morphological homoplasy.

Taxon 53: 415–428.

Raubeson, L. A. & Jansen, R. K. 1992. Chloroplast DNA evi-

dence on the ancient evolutionary split in vascular land

plants. Science 255: 1697–2699.

Reid, J. D., Plunkett, G. M. & Peters, G. A. 2006.

Phylogenetic relationships in the heterosporous fern genus

Azolla (Azollaceae) based on DNA sequence data from

three noncoding regions. Int. J. Pl. Sci. 167: 529–538.

Renzaglia, K. S., Duff, R. J., Nickrent, D. L. & Garbary, D.

J. 2000. Vegetative and reproductive innovations of early

land plants; implications for a unified phylogeny. Philos.

Trans., Ser. B 355: 769–793.

Rouhan, G., Dubuisson, J.-Y., Rakotondrainibe, F., Motley,

T. J., Mickel, J. T., Labat, J.-N. & Moran, R. C. 2004.

Molecular phylogeny of the fern genus Elaphoglossum

(Elaphoglossaceae) based on chloroplast non-coding DNA

sequences: contributions of species from the Indian Ocean

area. Molec. Phylog. Evol. 33: 745–763.

Sánchez-Baracaldo, P. 2004a. Phylogenetics and biogeogra-

phy of the neotropical fern genera Jamesonia and Erio-

sorus (Pteridaceae). Amer. J. Bot. 91: 274–284.

Sánchez-Baracaldo, P. 2004b. Phylogenetic relationships of

the subfamily Taenitoideae, Pteridaceae. Amer. Fern J. 94:

126–142.

Sano, R., Takamiya, M., Ito, M., Kurita, S. & Hasebe, M.

2000a. Phylogeny of the lady fern group, tribe

Physematieae (Dryopteridaceae), based on chloroplast

rbcL gene sequences. Molec. Phylog. Evol. 15: 403–413.

Sano, R., Takamiya, M., Kurita, S., Ito, M. & Hasebe, M.

2000b. Diplazium subsinuatum and D. tomitaroanum

should be moved to Deparia according to molecular, mor-

phological, and cytological characters. J. Pl. Res. 113:

157–163.

Schneider, H. 1996a. Vergleichende Wurzelanatomie der

Farne. Ph.D. dissertation. Univ. Zürich, Shaker, Aachen.

Schneider, H. 1996b. Root anatomy of Aspleniaceae and the

implications for systematics of the fern family. Fern Gaz.

12: 160–168.

Schneider, H., Janssen, T., Hovenkamp, P., Smith, A. R.,

Cranfill, R., Haufler, C. H. & Ranker, T. A. 2004a.

Phylogenetic relationships of the enigmatic Malesian fern

Thylacopteris (Polypodiaceae, Polypodiidae). Int. J. Pl.

Sci. 165: 1077–1087.

Schneider, H., Kreier, H.-P., Wilson, R. & Smith, A. R. 2006.

The Synammia enigma: evidence for a temperate lineage

of polygrammoid ferns (Polypodiaceae, Polypodiidae) in

southern South America. Syst. Bot. 31: 31–41.

Schneider, H. & Pryer, K. M. 2002. Structure and function of

spores in the aquatic heterosporous fern family

Marsileaceae. Int. J. Pl. Sci. 163: 485–505.

Schneider, H., Pryer, K. M., Cranfill, R., Smith, A. R. &

Wolf, P. G. 2002a. Evolution of vascular plant body plans:

a phylogenetic perspective. Pp. 330–364 in: Cronk, Q. C.

B., Bateman, R. M. & Hawkins, J. A. (eds.), Developmen-

tal Genetics and Plant Evolution. Taylor & Francis,

London.

Schneider, H., Ranker, T. A., Russell, S. J., Cranfill, R., Gei-

ger, J. M. O., Aguraiuja, R., Wood, K. R., Grundmann,

M., Kloberdanz, K. & Vogel, J. C. 2005. Origin of the

endemic fern genus Diellia coincides with the renewal of

Hawaiian terrestrial life in the Miocene. Proc. Roy. Soc.,

London, ser. B, Biol. Sci. 272: 455–460.

Schneider, H., Russell, S. J., Cox, C. J., Bakker, F., Hen-

derson, S., Gibby, M. & Vogel, J. C. 2004b. Chloroplast

phylogeny of asplenioid ferns based on rbcL and trnL-F

spacer sequences (Polypodiidae, Aspleniaceae) and its

implications for the biogeography. Syst. Bot. 29: 260–274.

Schneider, H., Schuettpelz, E., Pryer, K. M., Cranfill, R.,

Magallón, S. & Lupia, R. 2004c. Ferns diversified in the

shadow of angiosperms. Nature 428: 553–557.

Schneider, H., Smith, A. R., Cranfill, R., Haufler, C. H.,

Ranker, T. A. & Hildebrand, T. 2002b. Gymnogrammitis

dareiformis is a polygrammoid fern (Polypodiaceae)—

Resolving an apparent conflict between morphological

and molecular data. Pl. Syst. Evol. 234: 121–136.

Schneider, H., Smith, A. R., Cranfill, R., Hildebrand, T. E.,

Haufler, C. H. & Ranker, T. A. 2004d. Unraveling the

phylogeny of polygrammoid ferns (Polypodiaceae and

Grammitidaceae): exploring aspects of the diversification

of epiphytic plants. Molec. Phylog. Evol. 31: 1041–1063.

Schuettpelz, E., Korall, P. & Pryer, K. M. In press. Plastid

atpA data provide improved support for deep relationships

among ferns. Taxon 55.

Smith & al. • Fern classification55 (3) • August 2006: 705–731

723

Scotland, R. W. & Wortley, A. H. 2003. How many species of

seed plants are there? Taxon 52: 101–104.

Shmakov, A. I. 2003. Review of the family Woodsiaceae

(Diels) Herter of Eurasia. Pp. 49–64 in: Chandra, S. &

Srivastava, M. (eds.), Pteridology in the New Millennium.

Kluwer Academic Publishers, Dordrecht.

Skog, J. E., Mickel, J. T., Moran, R. C., Volovsek, M., Zim-

mer, E. A. 2004. Molecular studies of representative spe-

cies in the fern genus Elaphoglossum (Dryopteridaceae)

based on cpDNA sequences rbcL, trnL-F, and rps4-TRNS.

Int. J. Pl. Sci. 165: 1063–1075.

Skog, J. E., Mickel, J. T., Moran, R. & Zimmer, E. A. 2001.

Phylogeny of the fern genus Elaphoglossum based on two

chloroplast genes. P. 87 in: Botany 2001, Albuquerque,

New Mexico.New Mexico.

Skog, J. E., Zimmer, E. & Mickel, J. T. 2002. Additional sup-

port for two subgenera of Anemia (Schizaeaceae) from

data for the chloroplast intergenic spacer region trnL-F

and morphology. Amer. Fern J. 92: 119–130.

Smith, A. R. 1995 [1996]. Non-molecular phylogenetic hy-

potheses for ferns. Amer. Fern J. 85: 104–122.

Smith, A. R. & Cranfill, R. B. 2002. Intrafamilial relation-

ships of the thelypteroid ferns (Thelypteridaceae). Amer.

Fern J. 92: 131–149.

Smith, A. R., Kreier, H.-P., Haufler, C. H., Ranker, T. A. &

Schneider, H. In press. Serpocaulon (Polypodiaceae), a

new genus segregated from old Polypodium. Taxon 55.

Smith, A. R., Tuomisto, H., Pryer, K. M., Hunt, J. S. & Wolf,

P. G. 2001. Metaxya lanosa, a second species in the genus

and fern family Metaxyaceae. Syst. Bot. 26: 480–486.

Stein, D. B., Conant, D. S. & Valinski, A. E. C. 1996. The im-

plications of chloroplast DNA restriction site variation on

the classification and phylogeny of the Cyatheaceae. Pp.

235–254 in: Johns, R. J. (ed.), Holttum Memorial Volume.

Royal Botanic Gardens, Kew.

Stevenson, D. W. & Loconte, H. 1996. Ordinal and familial

relationships of pteridophyte genera. Pp. 435–467 in:

Camus, J. M., Gibby, M. & Johns, R. J. (eds.), Pteridology

in Perspective. Royal Botanic Gardens, Kew.

Sun, B.-Y., Kim, M. H., Kim, C. H. & Park, C.-W. 2001.

Mankyua (Ophioglossaceae): a new fern genus from

Cheju Island, Korea. Taxon 50: 1019–1024.

Tagawa, M. & Iwatsuki, K. 1972. Families and genera of the

pteridophytes known from Thailand. Mem. Fac. Sci. Kyoto

Univ., Ser. Biol. 5: 67–88.

Thorne, R. F. 2002. How many species of seed plants are

there? Taxon 51: 511–522.

Tindale, M. D. & Roy, S. K. 2002. A cytotaxonomic survey of

the Pteridophyta of Australia. Austr. Syst. Bot. 15:

839–937.

Tryon, A. F. 1964. Platyzoma—a Queensland fern with incip-

ient heterospory. Amer. J. Bot. 51: 939–942.

Tryon, A. F. & Lugardon, B. 1991. Spores of the Pteridophy-

ta. Springer-Verlag, New York.

Tryon, R. 1970. The classification of the Cyatheaceae. Contr.

Gray Herb. 200: 1–53.

Tryon, R. M. & Tryon, A. F. 1982. Ferns and Allied Plants,

with Special Reference to Tropical America. Springer-

Verlag, Berlin.

Tsutsumi, C. & Kato, M. 2005. Molecular phylogenetic study

on Davalliaceae. Fern Gaz. 17: 147–162.

Tsutsumi, C. & Kato, M. In press. Evolution of epiphytes in

Davalliaceae and related ferns. Bot. J. Linn. Soc.

van den Heede, C. J., Viane, R. L. L. & Chase, M. W. 2003.

Phylogenetic analysis of Asplenium subgenus Ceterach

(Pteridophyta: Aspleniaceae) based on plastid and nuclear

ribosomal ITS DNA sequences. Amer. J. Bot. 90: 481–493.

Vangerow, S., Teerkorn, T. & Knoop, V. 1999. Phylogenetic

information in the mitochondrial nad5 gene of pterido-

phytes: RNA editing and intron sequences. Pl. Biol. 1:

235–243.

Wagner, W. H., Jr. 1969. The construction of a classification.

Pp. 67–90 in: U. S. National Academy of Science,

Systematic Biology. U. S. Natl. Acad. Sci. Publ. No. 1692.

National Academy Press, Washington, D.C.

Wagner, W. H., Jr. & Smith, A. R. 1993. Pteridophytes of

North America. Pp. 247–266 in: Flora of North America

Editorial Committee (eds.), Flora of North America North

of Mexico, Vol. 1. Oxford Univ. Press, New York.

Wang, M.-L., Chen, Z.-D., Zhang, X.-C, Lu, S.-G. & Zhao,

G.-F. 2003. Phylogeny of the Athyriaceae: evidence from

chloroplast trnL-F region sequences. Acta Phytotax. Sin.

41: 416–426.

Wang, M.-L., Hsieh, Y.-T. & Zhao, G.-F. 2004. A revised sub-

division of the Athyriaceae. Acta Phytotax. Sin. 42:

524–527.

White, R. A. & Turner, M. D. 1988. Calochlaena, a new

genus of dicksonioid ferns. Amer. Fern J. 78: 86–95.

Wikström, N., Kenrick, P. & Vogel, J. C. 2002.

Schizaeaceae: a phylogenetic approach. Rev. Palaeobot.

Palynol. 119: 35–50.

Wikström, N. & Pryer, K. M. 2005. Incongruence between

primary sequence data and the distribution of a mitochon-

drial atp1 group II intron among ferns and horsetails.

Molec. Phylog. Evol. 36: 484–493.

Wolf, P. G. 1995 [1996]. Phylogenetic analyses of rbcL and

nuclear ribosomal RNA gene sequences in Dennstaedtia-

ceae. Amer. Fern J. 85: 306–327.

Wolf, P. G. 1996. Pteridophyte phylogenies based on analysis

of DNA sequences: a multiple gene approach. Pp.

203–215 in: Camus, J. M., Gibby, M. & Johns, R. J. (eds.),

Pteridology in Perspective. Royal Botanic Gardens, Kew.

Wolf, P. G. 1997. Evaluation of atpB nucleotide sequences for

phylogenetic studies of ferns and other pteridophytes.

Amer. J. Bot. 84: 1429–1440.

Wolf, P. G., Pryer, K. M., Smith, A. R. & Hasebe, M. 1998.

Phylogenetic studies of extant pteridophytes. Pp. 541–556

in: Soltis, D. E., Soltis, P. S., & Doyle, J. J. (eds.),

Molecular Systematics of Plants II. DNA Sequencing.

Kluwer Academic Publishers, Boston.

Wolf, P. G., Sipes, S. D., White, M. R., Martines, M. L.,

Pryer, K. M., Smith, A. R. & Ueda, K. 1999.

Phylogenetic relationships of the enigmatic fern families

Hymenophyllopsidaceae and Lophosoriaceae: evidence

from rbcL nucleotide sequences. Pl. Syst. Evol. 219:

263–270.

Wolf, P. G., Soltis, P. S. & Soltis, D. E. 1994. Phylogenetic

relationships of dennstaedtioid ferns: evidence from rbcL

sequences. Molec. Phylog. Evol. 3: 383–392.

Wu, S., Phan, K. L. & Xiang, J. 2005. A new genus and two

new species of ferns from Vietnam. Novon 15: 245–249.

Yatabe, Y., Nishida, H. & Murakami, N. 1999. Phylogeny of

Osmundaceae inferred from rbcL nucleotide sequences

and comparison to the fossil evidence. J. Pl. Res. 112:

Smith & al. • Fern classification 55 (3) • August 2006: 705–731

724

397–404.

Yatabe, Y., Watkins, J. E., Farrar, D. R. & Murakami, N.

2002. Genetic variation in populations of the morphologi-

cally and ecologically variable fern Stegnogramma pozoi

subsp. mollissima (Thelypteridaceae) in Japan. J. Pl. Res.

115: 29–38.

Zhang, G., Zhang, X. & Chen, Z. 2005. Phylogeny of cryp-

togrammoid ferns and related taxa based on rbcL

sequences. Nordic J. Bot. 23: 485–493.

Zhang, X.-C. & Nooteboom, H. P. 1998. A taxonomic revi-

sion of Plagiogyriaceae (Pteridophyta). Blumea 43:

401–469.

Smith & al. • Fern classification55 (3) • August 2006: 705–731

725

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

726

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

727

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

(“Antrophyaceae”).

•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

728

Appendix 4. Continued.

•Fam. Cystopteridaceae Schmakov, Turczaninowia 4: 60. 15 May 2001. – basionym: Cystopteridinae Payer, Bot. Crypt.: 198. 1850

(“Cystopterideae”).

•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,

1993).

•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

729

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

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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

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)

Micromorphology of fern spores as a tool in taxonomy of East-Central European species from the family Aspleniaceae (Polypodiopsida)

Article

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Jun 2024
ACTA SOC BOT POL

The family Aspleniaceae consists of ca. 730 fern species found primarily in the tropics but present also in the temperate zone. Recently, researchers have been increasingly interested in spore morphology, as it provides valuable information used in taxonomic and phylogenetic analyses. is study aimed to (1) describe in detail spore morphology in species of the family Aspleniaceae from Central and Eastern Europe; (2) attempt to use their features in taxonomic analyses; and (3) create a key to species identification on the basis of spores. e study was based on Asplenium spores of 10 species from 57 localities. e morphological type of spores was very similar in all the studied species, whereas spore microsculpture varied widely between and within species. is makes it difficult to use spores for species identification. Nevertheless, spore morphology allows the identification of six species, while the others form groups composed of 2–3 species each. In the group consisting of A. adulterinum, A. trichomanes, and A. ruta-muraria, the last species is distinguished by larger spore size (more than 50 μm long).

A new species of the equisetalean plant Equicalastrobus from the Middle Triassic of Argentina

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Jun 2024
ACTA PALAEONTOL POL

Deciphering the reticulate evolution and phylo-biogeography of Ant-Ferns Lecanopteris s.s

Preprint

Full-text available

Jun 2024

Southeast Asia is a biodiversity hotspot characterized by a complex paleogeography, and its Polypodiopsida flora is particularly diverse. While hybridization is recognized as common in ferns, investigations into the relationship between hybridization events and fern diversity are notably lacking. Lecanopteris s.s., an ant-associated fern, has been subject to debate regarding species delimitations primarily due to limited DNA markers and species sampling. Our study integrates 22 newly generated plastomes, 22 transcriptomes, and flow cytometry of all native species along with two cultivated hybrids. Our objective is to elucidate the reticulate evolutionary history within Lecanopteris through the integration of phylo-biogeography reconstruction, gene flow inference, and genome size estimation. Key findings of our study include: (1) An enlarged plastome size in Lecanopteris, attributed to extreme expansion of the Inverted Repeat (IR) regions; (2) The traditional 'pumila' and 'crustacea' groups are paraphyletic; (3) Significant cytonuclear discordance attributed to gene flow; (4) Natural hybridization and introgression in the 'pumila' and 'darnaedii' groups; (5) L. luzonensis is the maternal parent of L. 'Yellow Tip', with L. pumila suggested as a possible paternal parent; (6) L. 'Tatsuta' is a hybrid between L. luzonensis and L. crustacea; (7) Lecanopteris first diverged during the Neogene and then during the middle Miocene climatic optimum in Indochina and Sundaic regions. In conclusion, the biogeographic history and speciation of Lecanopteris have been profoundly shaped by past climate changes and geodynamics of Southeast Asia. Dispersals, hybridization and introgression between species act as pivotal factors in the evolutionary trajectory of Lecanopteris.

PlantSR: Super-Resolution Improves Object Detection in Plant Images

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Jun 2024

Recent advancements in computer vision, especially deep learning models, have shown considerable promise in tasks related to plant image object detection. However, the efficiency of these deep learning models heavily relies on input image quality, with low-resolution images significantly hindering model performance. Therefore, reconstructing high-quality images through specific techniques will help extract features from plant images, thus improving model performance. In this study, we explored the value of super-resolution technology for improving object detection model performance on plant images. Firstly, we built a comprehensive dataset comprising 1030 high-resolution plant images, named the PlantSR dataset. Subsequently, we developed a super-resolution model using the PlantSR dataset and benchmarked it against several state-of-the-art models designed for general image super-resolution tasks. Our proposed model demonstrated superior performance on the PlantSR dataset, indicating its efficacy in enhancing the super-resolution of plant images. Furthermore, we explored the effect of super-resolution on two specific object detection tasks: apple counting and soybean seed counting. By incorporating super-resolution as a pre-processing step, we observed a significant reduction in mean absolute error. Specifically, with the YOLOv7 model employed for apple counting, the mean absolute error decreased from 13.085 to 5.71. Similarly, with the P2PNet-Soy model utilized for soybean seed counting, the mean absolute error decreased from 19.159 to 15.085. These findings underscore the substantial potential of super-resolution technology in improving the performance of object detection models for accurately detecting and counting specific plants from images. The source codes and associated datasets related to this study are available at Github.

The Relationship of Terrestrial Fern Species (Pteridophyta) at Salodik Waterfall Central Sulawesi Based on Morphological Characteristics

Article

Full-text available

May 2024

Pteridophyta ferns are a group of vascular plants that have a very long evolutionary history. This study aims to describe the relationship of terrestrial fern plant species (Pteridophyta) at Salodik Waterfall Central Sulawesi based on morphological characteristics. The method used was exploration method with the sampling technique being free collection. The data were processed using Microsoft Excel 2013 and then subjected to cluster analysis using the UPGMA (Unweighted Pair-Group Method with Arithmetic Average) method in the PAST 4.03 software (Paleontological Statistics). The research results obtained a total of 13 terrestrial fern plant species, namely Asplenium nidus, Diplazium esculentum, Davallia denticulata, Lindsaea scandens var. terrestris, Nephrolepis biserrata, Angiopteris evecta, Phymatosorus longissima, Pteris biaurita, Pteris fauriei, Selaginella doederleinii, Chingia clavipilosa, Macrothelypteris torresiana and Sphaerostephanos heterocarpus. The dendrogram of kinship between 13 species of terrestrial ferns is formed in 2 main clusters with 4 relationships categories, namely very near, near, not near, and not very near. The first category of relationship is very near with the highest Similarity Index (IS) value, namely between Macrothelypteris torresiana with Davalia denticulata (IS=86.5) and Lindsaea scandens with Pteris faurei (IS=75.2). The second category of near relationship is between Pteris biaurita with node 3 (IS=68.2), Selaginella doederleinii with node 4 (IS=64), and Diplazium esculentum with Chingia clavipilosa (IS=55.6). The third category of not near relationship is between Nephrolepis biserrata with Phymatosorus longissima (IS=27.3). The fourth category of not very near relationship with the lowest Similarity Index (IS) value is between Angiopteris evecta with Sphaerostephanos heterocarpus (IS=22.8) and Asplenium nidus with sub division Ia (IS=21.2).

Asplenium sylvaticum (Aspleniaceae), a new fern species from Yungas and Paraná forests, Neotropical region

Article

Full-text available

Jun 2024
AN ACAD BRAS CIENC

A new species of Asplenium L. (Aspleniaceae) is described and illustrated. Asplenium sylvaticum is endemic of the Yungas and Paraná biogeographic provinces in the Southern Cone. The main diagnostic characters for this taxon are: apical pinnae with one to four caudate lobes at the base, similar in size to lateral pinnae, smooth pinnate blades with 18-36 pairs of caudate pinnae and spores with reticulate perispore, winged folds with scarce equinulaes on the margins and the laesurae wide with smooth margin. The new species belongs to the A. serra species complex group, characterised by xeromorphic habit, creeping rhizome and coriaceous fronds with branched scales on both surfaces, and resembles the other species inhabiting in Argentina, A. achalense and A. serra, in overall leaf architecture and shape but differs of the aforementioned species (and the other species of the A. serra complex) by the unique combination of characters of its rhizomatic scales, fronds and spores. A key with the morphological and palynological characters that differentiate A. sylvaticum from the other taxa belonging to the A. serra complex in the South American Cone and Bolivia is given.

Maiaspora: the hallmark of gleichenioid ferns (Gleicheniales) from the early Carboniferous

Article

Jun 2024

Dear colleagues, you can download our paper using my personal link. Thank you for the interest to our research ------https://onlinelibrary.wiley.com/share/author/G997NYRUBFZHTTWYEFPE?target=10.1002/spp2.1561 ------------------------------------------------------------ The Palaeozoic origin of gleichenioid ferns (Gleicheniales) has aroused curiosity for almost a century. The oldest definite sign of this ancient clade is the spore wall ultrastructure of Radiitheca dobranyana from the Moscovian of central Europe. However, familial assignment of this fern fossil is uncertain. A comparison of its in situ spores with those of the dispersed genus Maiaspora, from the Visean, suggests that they may represent the same group of leptosporangiates. Although these spores have a common metareticuloid sculpture, the ultra-thin structure of the dispersed counterparts remained unknown. Study of the ultrastructure of Maiaspora concava and M. panopta from central Russia and Northumberland (UK) using transmission electron microscopy shows that the wall of Maiaspora has a middle exospore layer composed of foliated lamellae. These spread from the base of the aperture slit to the subequatorial margins of kyrtome areas. The lamellae do not overlap the aperture slit and terminate at between two-thirds and three-quarters of the slit height. The miospores have a three-layered exospore in the interradial areas, while non-apertural parts of the exospore consist of two layers. Such an ultrastructure is shared by extant Gleicheniales and fossil R. dobranyana. The combined evidence from the morphology, ultrastructure and spatiotemporal distribution of Maiaspora, as well as radioisotope dating and additional plant macrofossils, suggests that the origin of the Gleicheniales stem was related to closure of the Rheic Ocean. Apparently, the first Maiaspora-producing gleichenioids emerged as neoendemic terrestrial ferns on the peri-Gondwanan volcanic arcs.

Samambaias e licófitas de um mosaico vegetacional do bioma Pampa, Estado do Rio Grande do Sul, Brasil

Article

Full-text available

Jun 2024

O domínio Pampeano constitui um mosaico vegetacional de campo-floresta, que abriga uma alta riqueza de espécies, incluindo as plantas vasculares dispersas por esporos. O objetivo deste estudo foi inventariar as samambaias e licófitas em um fragmento do Pampa, localizado na Fazenda Estância da Quinta, no município de Rio Pardo, Estado do Rio Grande do Sul. A amostragem ocorreu através do método de caminhamento expedito, realizada em 15 viagens de campo. O material coletado foi identificado através de bibliografia especializada, com posterior incorporação no Herbário da Universidade de Caxias do Sul (HUCS). Atributos ecológicos, geográficos e de conservação das espécies foram registrados em campo e através de busca bibliográfica. O inventário resultou no reconhecimento de 46 espécies de samambaias e uma licófita (Isoetes weberi Herter), pertencentes a 13 famílias. O hábito terrestre (51%) foi predominante, bem como a ocorrência preferencial no interior dos fragmentos florestais (66%). Duas espécies constam em listas de espécies ameaçadas de extinção. Nossos dados reforçam a importância da preservação do mosaico paisagístico do Pampa para manutenção da diversidade deste grupo de plantas do RS.

Morphology and ultrastructure of cicatricose spores found in an isolated sporangium-bearing structure of a schizaealean fern from the Lower Cretaceous rocks at Casal do Borracho, Torres Vedras, western Portugal

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Jun 2024
REV PALAEOBOT PALYNO

The Early Cretaceous tree fern Acanthopteris (Dicksoniaceae): New insight into fossil records, species diversity, palaeogeography and palaeoclimate

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CRETACEOUS RES

Palaeobiological and phylogenetic implications of anatomically-preserved archaeocalamites from the Dinantian of Oxroad Bay and Loch Humphrey Burn, southern Scotland.

Article

Full-text available

Jan 1991
Palaeontograph Abteilung B

Richard Bateman

The Radiation of Vascular Plants

Chapter

Jul 2004

This edited volume is provides an authoritative synthesis of knowledge about the history of life. All the major groups of organisms are treated, by the leading workers in their fields. With sections on: The Importance of Knowing the Tree of Life; The Origin and Radiation of Life on Earth; The Relationships of Green Plants; The Relationships of Fungi; and The Relationships of Animals. This book should prove indispensable for evolutionary biologists, taxonomists, ecologists interested in biodiversity, and as a baseline sourcebook for organismic biologists, botanists, and microbiologists. An essential reference in this fundamental area.

PLATYZOMA-A QUEENSLAND FERN WITH INCIPIENT HETEROSPORY

Article

Oct 1964

Alice F. Tryon

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.

Pteridophytes and Gymnosperms

Book

Jan 1990

Cheilanthoid Ferns (Pteridaceae: Cheilanthoideae) in the Southwestern United States and Adjacent Mexico-A Molecular Phylogenetic Reassessment of Generic Lines

Article

Jan 1998

Spores of the Pteridophyta

Book

Jan 1991

Ferns and Allied Plants

Book

Jan 1982

The Chinese fern families and genera: Systematic arrangement and historical origin

Article

Jan 1978
ACTA PHYTOTAXON SIN

R.C. Ching

PTERIDOPHYTA

Chapter

Dec 1959

RODOLFO E.G. PICHI-SERMOLLI

Review of the Family Woodsiaceae (Diels) Herter of Eurasia

Chapter

Jan 2003

Alexandr Ivanovich Shmakov

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.

A Classification for Extant Ferns

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