A pteridophyte is a
vascular plant
Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
(with
xylem
Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
and
phloem
Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
) that reproduces by means of
spore
In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s. Because pteridophytes produce neither
flower
Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s nor
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s, they are sometimes referred to as "
cryptogams", meaning that their means of reproduction is hidden. They are also the ancestors of the plants we see today.
Fern
The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s,
horsetail
''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds.
''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which ...
s (often treated as ferns), and
lycophytes (
clubmoss
Lycopodiopsida is a class of vascular plants also known as lycopsids, lycopods, or lycophytes. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching stems bearing simple leaves ...
es,
spikemosses, and
quillworts) are all pteridophytes. However, they do not form a
monophyletic group because ferns (and horsetails) are more closely related to
seed plant
A seed plant or spermatophyte (; New Latin ''spermat-'' and Greek ' (phytón), plant), also known as a phanerogam (taxon Phanerogamae) or a phaenogam (taxon Phaenogamae), is any plant that produces seeds. It is a category of embryophyte (i.e. la ...
s than to lycophytes. "Pteridophyta" is thus no longer a widely accepted taxon, but the term ''pteridophyte'' remains in common parlance, as do ''pteridology'' and ''pteridologist'' as a science and its practitioner, for example by the International Association of Pteridologists and the
Pteridophyte Phylogeny Group
The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
.
Etymology
The name ''Pteridophyte'' is a
Neo-Latin
Neo-LatinSidwell, Keith ''Classical Latin-Medieval Latin-Neo Latin'' in ; others, throughout. (also known as New Latin and Modern Latin) is the style of written Latin used in original literary, scholarly, and scientific works, first in Italy d ...
compound word created by English speakers around 1880. It is formed from the prefix ''pterido-'' meaning fern, a Latin borrowing of the Greek word ''pterís'' which derives from ''pterón'' meaning feather. The suffix, ''-phyte'', is a suffix meaning plant from the ancient Greek word phyton (φυτόν).
Description
Pteridophytes (ferns and lycophytes) are free-sporing
vascular plant
Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s that have a
life cycle with
alternating, free-living
gametophyte
A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
and
sporophyte
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
phases that are independent at maturity. The body of the sporophyte is well differentiated into roots, stem and leaves. The root system is always
adventitious
Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant ...
. The stem is either underground or aerial. The leaves may be
microphylls or megaphylls. Their other common characteristics include vascular plant
apomorphies
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
(e.g.,
vascular tissue
Vascular tissue is a complex transporting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. T ...
) and
land plant
The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophyt ...
plesiomorphies (e.g.,
spore
In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
dispersal and the absence of
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s).
Taxonomy
Phylogeny
Of the pteridophytes, ferns account for nearly 90% of the extant diversity. Smith et al. (2006), the first higher-level pteridophyte classification published in the
molecular phylogenetic
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
era, considered the ferns as monilophytes, as follows:
*
Division Tracheophyta
Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
(tracheophytes) - vascular plants
** Subdivision
Lycopodiophyta (lycophytes) - less than 1% of extant vascular plants
**
Sub division Euphyllophytina (euphyllophytes)
*** Infradivision
Moniliformopses (monilophytes)
*** Infradivision
Spermatophyta
A seed plant or spermatophyte (; New Latin ''spermat-'' and Greek ' (phytón), plant), also known as a phanerogam (taxon Phanerogamae) or a phaenogam (taxon Phaenogamae), is any plant that produces seeds. It is a category of embryophyte (i.e. l ...
- seed plants, ~260,000 species
where the monilophytes comprise about 9,000 species, including
horsetails (
Equisetaceae),
whisk ferns (Psilotaceae), and all
eusporangiate and all
leptosporangiate ferns. Historically both lycophytes and monilophytes were grouped together as pteridophytes (ferns and fern allies) on the basis of being spore-bearing ("seed-free"). In Smith's molecular phylogenetic study the ferns are characterised by
lateral root origin in the
endodermis
The endodermis is the innermost layer of cortex in land plants. It is a cylinder of compact living cells, the radial walls of which are impregnated with hydrophobic substances ( Casparian strip) to restrict apoplastic flow of water to the inside ...
, usually
mesarch protoxylem in shoots, a pseudoendospore,
plasmodial tapetum, and
sperm cells with 30-1000
flagella
A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
. The term "moniliform" as in Moniliformopses and monilophytes means "bead-shaped" and was introduced by Kenrick and Crane (1997) as a scientific replacement for "fern" (including Equisetaceae) and became established by Pryer et al. (2004). Christenhusz and Chase (2014) in their review of classification schemes provide a critique of this usage, which they discouraged as irrational. In fact the alternative name
Filicopsida was already in use. By comparison "lycopod" or lycophyte (club moss) means wolf-plant. The term "
fern ally" included under Pteridophyta generally refers to vascular spore-bearing plants that are not ferns, including lycopods, horsetails, whisk ferns and water ferns (
Marsileaceae
Marsileaceae () is a small family of heterosporous aquatic and semi-aquatic ferns, though at first sight they do not physically resemble other ferns. The group is commonly known as the "pepperwort family" or as the "water-clover family" becau ...
,
Salviniaceae and ''
Ceratopteris''). This is not a natural grouping but rather a convenient term for non-fern, and is also discouraged, as is eusporangiate for non-leptosporangiate ferns.
However both Infradivision and Moniliformopses are also invalid names under the
International Code of Botanical Nomenclature
The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN or ICNafp) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all tho ...
. Ferns, despite forming a
monophyletic
In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria:
# the grouping contains its own most recent co ...
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, are formally only considered as four
classes (
Psilotopsida;
Equisetopsida;
Marattiopsida;
Polypodiopsida
The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
), 11
orders
Order, ORDER or Orders may refer to:
* A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* H ...
and 37
families
Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
, without assigning a higher
taxonomic rank
In biology, taxonomic rank (which some authors prefer to call nomenclatural rank because ranking is part of nomenclature rather than taxonomy proper, according to some definitions of these terms) is the relative or absolute level of a group of or ...
.
Furthermore, within the Polypodiopsida, the largest grouping, a number of informal clades were recognised, including leptosporangiates, core leptosporangiates,
polypods (Polypodiales), and eupolypods (including
Eupolypods I
Polypodiineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods I in earlier systems, and to the very broadly defined family Polypodiaceae in the classification of Christenhusz & Chase (2014). It probably ...
and
Eupolypods II).
In 2014
Christenhusz and
Chase, summarising the known knowledge at that time, treated this group as two separate unrelated taxa in a consensus classification;
*
Lycopodiophyta (lycopods) 1 subclass, 3 orders, each with one family, 5 genera, approx. 1,300 species
*
Polypodiophyta (ferns) 4 subclasses, 11 orders, 21 families, approx. 212 genera, approx. 10,535 species
** Subclass
Equisetidae Warm.
** Subclass
Ophioglossidae
Ophioglossidae is one of the four subclasses of Polypodiopsida (ferns). This subclass consists of the ferns commonly known as whisk ferns, grape ferns, adder's-tongues and moonworts. It is equivalent to the class Psilotopsida in previous treat ...
Klinge
** Subclass
Marattiidae
Marattiaceae is the only family of extant (living) ferns in the order Marattiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), Marattiales is the only order in the subclass Marattiidae. The family has six genera and about ...
Klinge
** Subclass
Polypodiidae Cronquist, Takht. & Zimmerm.
These subclasses correspond to Smith's four classes, with Ophioglossidae corresponding to Psilotopsida.
The two major groups previously included in Pteridophyta are
phylogenetically
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
related as follows:
Subdivision
Pteridophytes consist of two separate but related classes, whose nomenclature has varied. The system put forward by the Pteridophyte Phylogeny Group in 2016,
PPG I, is:
* Class
Lycopodiopsida
Lycopodiopsida is a class of vascular plants also known as lycopsids, lycopods, or lycophytes. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching stems bearing simple leaves ...
Bartl. – lycophytes: clubmosses, quillworts and spikemosses; 3 extant orders
::*Order
Lycopodiales DC. ex Bercht. & J.Presl – clubmosses; 1 extant family
::*Order
Isoetales
Isoetales, sometimes also written Isoëtales, is an order of plants in the class Lycopodiopsida.
There are about 140-150 living species, all of which are classified in the genus '' Isoetes'' (quillworts), with a cosmopolitan distribution, but ...
Prantl – quillworts; 1 extant family
::*Order
Selaginellales Prantl – spikemosses; 1 extant family
* Class
Polypodiopsida
The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
Cronquist, Takht. & W.Zimm. – ferns; 11 extant orders
** Subclass
Equisetidae Warm. – horsetails; 1 extant order, family and genus (''
Equisetum
''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds.
''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which ...
'')
*** Order
Equisetales DC. ex Bercht. & J.Presl – 1 extant family
** Subclass
Ophioglossidae
Ophioglossidae is one of the four subclasses of Polypodiopsida (ferns). This subclass consists of the ferns commonly known as whisk ferns, grape ferns, adder's-tongues and moonworts. It is equivalent to the class Psilotopsida in previous treat ...
Klinge – 2 extant orders
*** Order
Psilotales Prant – whisk ferns; 1 extant family
*** Order
Ophioglossales Link – grape ferns; 1 extant family
** Subclass
Marattiidae
Marattiaceae is the only family of extant (living) ferns in the order Marattiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), Marattiales is the only order in the subclass Marattiidae. The family has six genera and about ...
Klinge – marattioid ferns; 1 extant order
*** Order
Marattiales
Marattiaceae is the only family of extant (living) ferns in the order Marattiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), Marattiales is the only order in the subclass Marattiidae. The family has six genera and about ...
Link – 1 extant family
** Subclass
Polypodiidae Cronquist, Takht. & W.Zimm. – leptosporangiate ferns; 7 extant orders
*** Order
Osmundales
Osmundaceae (royal fern family) is a family of ferns containing four to six extant genera and 18–25 known species. It is the only living family of the order Osmundales in the class Polypodiopsida (ferns) or in some classifications the only or ...
Link – 1 extant family
*** Order
Hymenophyllales
The Hymenophyllaceae, the filmy ferns and bristle ferns, are a family of two to nine genera (depending on classification system) and about 650 known species of ferns, with a subcosmopolitan distribution, but generally restricted to very damp pla ...
A.B.Frank – 1 extant family
*** Order
Gleicheniales
Gleicheniales is an order of ferns in the subclass Polypodiidae (the leptosporangiate ferns). The Gleicheniales has spore records potentially as early as the Early Carboniferous, but the oldest unambiguous macrofossil records date to the Early ...
Schimp – 3 extant families
*** Order
Schizaeales Schimp. – 3 extant families
*** Order
Salviniales
The order Salviniales (formerly known as the Hydropteridales and including the former Marsileales) is an order of ferns in the class Polypodiopsida.
Description
Salviniales are all aquatic and differ from all other ferns in being heterosporous, ...
Link – 2 extant families
*** Order
Cyatheales
The order Cyatheales, which includes most tree ferns, is a Taxonomy (biology), taxonomic order of the fern class, Polypodiopsida. No clear morphology (biology), morphological features characterize all of the Cyatheales, but DNA sequence data indi ...
A.B.Frank – 8 extant families
*** Order
Polypodiales
The Order (biology), order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including Tropics, tropical, semitropical and Temperate clima ...
Link – 26 extant families
In addition to these living groups, several groups of pteridophytes are now
extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
and known only from
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s. These groups include the
Rhyniopsida,
Zosterophyllopsida,
Trimerophytopsida, the
Lepidodendrales and the
Progymnospermopsida.
Modern studies of the land plants agree that seed plants emerged from
pteridophytes more closer to ferns than lycophytes. Therefore, pteridophytes do not form a clade but constitute a
paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
grade.
Life cycle

Just as with
bryophyte
Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s and
spermatophyte
A seed plant or spermatophyte (; New Latin ''spermat-'' and Greek ' (phytón), plant), also known as a phanerogam (taxon Phanerogamae) or a phaenogam (taxon Phaenogamae), is any plant that produces seeds. It is a category of embryophyte (i.e. la ...
s (seed plants), the life cycle of pteridophytes involves
alternation of generations
Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploi ...
. This means that a
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
generation (the sporophyte, which produces spores) is followed by a
haploid
Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
generation (the gametophyte or
prothallus
A prothallus, or prothallium, (from Latin ''pro'' = forwards and Greek ''θαλλος'' (''thallos'') = twig) is usually the gametophyte stage in the life of a fern or other pteridophyte. Occasionally the term is also used to describe the young ...
, which produces
gametes
A gamete ( ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. The name gamete was introduced by the Ge ...
). Pteridophytes differ from bryophytes in that the sporophyte is branched and generally much larger and more conspicuous, and from seed plants in that both generations are independent and free-living. The sexuality of pteridophyte gametophytes can be classified as follows:
*
Dioicous
Dioicy () is a sexual system in non-vascular plants where archegonia (female organs) and antheridia (male organs) are produced on separate plants in the gametophyte phase. It is one of the two main sexual systems in bryophytes, the other being ...
: each individual gametophyte is either male (producing
antheridia and hence
sperm
Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
) or female (producing
archegonia and hence
egg cell
The egg cell or ovum (: ova) is the female Reproduction, reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female game ...
s).
*
Monoicous
Monoicy () is a sexual system in haploid plants (mainly bryophytes) where both sperm and eggs are produced on the same gametophyte, in contrast with dioicy, where each gametophyte produces only sperm or eggs but never both.Crandall-Stotler, B.J ...
: each individual gametophyte produces both antheridia and archegonia and can function both as a male and as a female.
*:
Protandrous: the antheridia mature before the archegonia (male first, then female).
*:
Protogynous: the archegonia mature before the antheridia (female first, then male).
These terms are ''not'' the same as
monoecious
Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
and
dioecious
Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, which refer to whether a seed plant's sporophyte bears both male and female gametophytes, i.e., produces both pollen and seeds, or just one.
See also
*
Embryophyte
The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...
* Fern ally
* Plant sexuality
References
Bibliography
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External links
British Pteridological SocietyAnnual Review of Pteridological Research *
Pteridophytes Test Questions for Papers And Quizzes [Important/nowiki>]
{{Authority control
Cryptogams
Plants
Paraphyletic groups