These plants are generally
sporophyte-oriented; that is, the normal plant is the
diploid sporophyte, with the only
haploid structure being the
gametophyte (
prothallium) in season. This basic pattern is like that found in the seed plants but with an important exception. Unlike the seed plants, the pteridophytes have a gametophyte stage that is free-living. As a result, pteridophyte sexuality is more complicated than that of the seed plants. There are several basic categories of sexuality in pteridophytes. The terms distinguish between types of gametophyte sexuality:
Dioicous pteridophytes produce only
antheridia (male organs) or
archegonia (female organ) on a single gametophyte body.
Monoicous pteridophytes produce both
antheridia and
archegonia on the same gametophyte body.
Protandrous pteridophytes produce the male antheridia first, and then their female archegonia.
Protogynous pteridophytes produce the archegonia first, followed by the antheridia.
Notice that these terms are not the same as monoecious and dioecious, which refer to whether or not a
sporophyte plant bears one or both kinds of gametophyte. Those terms apply only to
seed plants.These plants are generally
sporophyte-oriented; that is, the normal plant is the
diploid sporophyte, with the only
haploid structure being the
gametophyte (
prothallium) in season. This basic pattern is like that found in the seed plants but with an important exception. Unlike the seed plants, the pteridophytes have a gametophyte stage that is free-living. As a result, pteridophyte sexuality is more complicated than that of the seed plants. There are several basic categories of sexuality in pteridophytes. The terms distinguish between types of gametophyte sexuality:
Dioicous pteridophytes produce only
antheridia (male organs) or
archegonia (female organ) on a single gametophyte body.
Monoicous pteridophytes produce both
antheridia and
archegonia on the same gametophyte body.
Protandrous pteridophytes produce the male antheridia first, and then their female archegonia.
Protogynous pteridophytes produce the archegonia first, followed by the antheridia.
Notice that these terms are not the same as monoecious and dioecious, which refer to whether or not a
sporophyte plant bears one or both kinds of gametophyte. Those terms apply only to
seed plants.Ferns first appear in the fossil record in the early-
Carboniferous period. By the
Triassic, the first evidence of ferns related to several modern families appeared. The "great fern radiation" occurred in the late-
Cretaceous, when many modern families of ferns first appeared.
Ferns have traditionally been grouped in the Class Filices, but modern classifications assign them their own division in the plant kingdom, called Pteridophyta.
Traditionally, three discrete groups of plants have been considered ferns: two groups of eusporangiate ferns — families
Ophioglossaceae (
adders-tongues,
moonworts, and
grape-ferns) and
Marattiaceae — and the
leptosporangiate ferns. The Marattiaceae are a primitive group of tropical ferns with a large, fleshy rhizome, and are now thought to be a sibling taxon to the main group of ferns, the leptosporangiate ferns. Several other groups of plants were considered "
fern allies": the
clubmosses,
spikemosses, and
quillworts in the
Lycopodiophyta, the
whisk ferns in
Psilotaceae, and the
horsetails in the
Equisetaceae. More recent genetic studies have shown that the Lycopodiophyta are only distantly related to any other
vascular plants, having radiated evolutionarily at the base of the vascular plant
clade, while both the whisk ferns and horsetails are as much "true" ferns as are the Ophioglossoids and Marattiaceae. In fact, the whisk ferns and Ophioglossoids are demonstrably a clade, and the horsetails and Marattiaceae are arguably another clade. Molecular data — which remain poorly constrained for many parts of the plants' phylogeny — have been supplemented by recent morphological observations supporting the inclusion of Equisetaceae within the ferns, notably relating to the construction of their sperm, and peculiarities of their roots (Smith et al 2006, and references therein).
One possible means of treating this situation is to consider only the leptosporangiate ferns as "true" ferns, while considering the other three groups as "fern allies". In practice, numerous classification schemes have been proposed for ferns and fern allies, and there has been little consensus among them. A new classification by Smith et al. (2006) is based on recent molecular systematic studies, in addition to morphological data. This classification divides ferns into four classes:
Psilotopsida
Equisetopsida
Marattiopsida
Polypodiopsida
The last group includes most plants familiarly known as ferns. Modern research supports older ideas based on morphology that the Osmundaceae diverged early in the evolutionary history of the leptosporangiate ferns; in certain ways this family is intermediate between the eusporangiate ferns and the leptosporangiate ferns.
The complete classification scheme proposed by Smith et al. (2006; alternative names in brackets):
Class
PsilotopsidaOrder
OphioglossalesFamily
Ophioglossaceae (incl.
Botrychiaceae,
Helminthostachyaceae)
Order
PsilotalesFamily
Psilotaceae (incl.
Tmesipteridaceae)
Class
Equisetopsida [=Sphenopsida]
Order
EquisetalesFamily
EquisetaceaeClass
MarattiopsidaOrder
MarattialesFamily
Marattiaceae (incl.
Angiopteridaceae,
Christenseniaceae,
Danaeaceae,
Kaulfussiaceae)
Class
Pteridopsida [=Filicopsida, Polypodiopsida]
Order
OsmundalesFamily
OsmundaceaeOrder
HymenophyllalesFamily
Hymenophyllaceae (incl.
Trichomanaceae)
Order
GleichenialesFamily
Gleicheniaceae (incl.
Dicranopteridaceae,
Stromatopteridaceae)
Family
Dipteridaceae (incl.
Cheiropleuriaceae)
Family
MatoniaceaeOrder
SchizaealesFamily
LygodiaceaeFamily
Anemiaceae (incl.
Mohriaceae)
Family
SchizaeaceaeOrder
SalvinialesFamily
Marsileaceae (incl.
Pilulariaceae)
Family
Salviniaceae (incl.
Azollaceae)
Order
CyathealesFamily
ThyrsopteridaceaeFamily
LoxomataceaeFamily
CulcitaceaeFamily
PlagiogyriaceaeFamily
CibotiaceaeFamily
Cyatheaceae (incl.
Alsophilaceae,
Hymenophyllopsidaceae)
Family
Dicksoniaceae (incl.
Lophosoriaceae)
Family
MetaxyaceaeOrder
PolypodialesFamily
Lindsaeaceae (incl.
Cystodiaceae,
Lonchitidaceae)
Family
SaccolomataceaeFamily
Dennstaedtiaceae (incl.
Hypolepidaceae,
Monachosoraceae,
Pteridiaceae)
Family
Pteridaceae (incl.
Acrostichaceae,
Actiniopteridaceae,
Adiantaceae,
Anopteraceae,
Antrophyaceae,
Ceratopteridaceae,
Cheilanthaceae,
Cryptogrammaceae,
Hemionitidaceae,
Negripteridaceae,
Parkeriaceae,
Platyzomataceae,
Sinopteridaceae,
Taenitidaceae,
Vittariaceae)
Family
AspleniaceaeFamily
ThelypteridaceaeFamily
Woodsiaceae (incl.
Athyriaceae,
Cystopteridaceae)
Family
Blechnaceae (incl.
Stenochlaenaceae)
Family
OnocleaceaeFamily
Dryopteridaceae (incl.
Aspidiaceae,
Bolbitidaceae,
Elaphoglossaceae,
Hypodematiaceae,
Peranemataceae)
Family
OleandraceaeFamily
DavalliaceaeFamily
Polypodiaceae (incl.
Drynariaceae,
Grammitidaceae,
Gymnogrammitidaceae,
Loxogrammaceae,
Platyceriaceae,
Pleurisoriopsidaceae)