Lycopsid joggins mcr1.JPG
   HOME

TheInfoList



OR:

Lycopodiopsida is a class of
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching stems bearing simple leaves called
microphylls In plant anatomy and evolution a microphyll (or lycophyll) is a type of plant leaf with one single, unbranched leaf vein. Plants with microphyll leaves occur early in the fossil record, and few such plants exist today. In the classical concept of ...
and reproduce by means of spores borne in
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cyc ...
on the sides of the stems at the bases of the leaves. Although living species are small, during the Carboniferous, extinct tree-like forms formed huge forests that dominated the landscape and contributed to
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when ...
deposits. The nomenclature and classification of plants with microphylls varies substantially among authors. A consensus classification for extant (living) species was produced in 2016 by the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relatio ...
(PPG I), which places them all in the class Lycopodiopsida, which includes the classes Isoetopsida and Selaginellopsida used in other systems. (See Table 2.) Alternative classification systems have used ranks from division (phylum) to subclass. In the PPG I system, the class is divided into three orders,
Lycopodiales The Lycopodiaceae (class Lycopodiopsida, order Lycopodiales) are an old family of vascular plants, including all of the core clubmosses and firmosses, comprising 16 accepted genera and about 400 known species. This family originated about 380 mil ...
,
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 ...
and
Selaginellales ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
.


Characteristics

Club-mosses (Lycopodiales) are homosporous, but the genera ''
Selaginella ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
'' (spikemosses) and ''
Isoetes ''Isoetes'', commonly known as the quillworts, is the only extant genus of plants in the family Isoetaceae, which is in the class of lycopods. There are currently 192 recognized species, with a cosmopolitan distribution but with the individual ...
'' (quillworts) are heterosporous, with female spores larger than the male, and gametophytes forming entirely within the spore walls. Which makes the club-moss gametophyte monoecious (both male and female sex organs forming on the same gametophyte) and the gametophyte of spikemosses and quillworts dioecious (separate male and female). Only the female gametophyte produces sporophytes. A few species of ''Selaginella'' such as '' S. apoda'' and '' S. rupestris'' are also
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the ...
; the gametophyte develops on the mother plant, and only when the sporophyte's primary shoot and root is developed enough for independence is the new plant dropped to the ground. Club-moss
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 ...
s are
mycoheterotrophic Myco-heterotrophy (from Greek μύκης , "fungus", ἕτερος ', "another", "different" and τροφή ', "nutrition") is a symbiotic relationship between certain kinds of plants and fungi, in which the plant gets all or part of its food fro ...
and long-lived, residing underground for several years before emerging from the ground and progressing to the sporophyte stage.


Taxonomy


Phylogeny

The extant lycophytes are
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s (tracheophytes) with microphyllous leaves, distinguishing them from the
euphyllophyte The euphyllophytes are a clade of plants within the tracheophytes (the vascular plants). The group may be treated as an unranked clade, a division Division or divider may refer to: Mathematics *Division (mathematics), the inverse of multipli ...
s (plants with megaphyllous leaves). The sister group of the extant lycophytes and their closest extinct relatives are generally believed to be the
zosterophyll The zosterophylls are a group of extinct land plants that first appeared in the Silurian period. The taxon was first established by Banks in 1968 as the subdivision Zosterophyllophytina; they have since also been treated as the division Zoste ...
s, a paraphyletic or
plesion In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
group. Ignoring some smaller extinct taxa, the evolutionary relationships are as shown below. , there was broad agreement, supported by both molecular and morphological evidence, that the extant lycophytes fell into three groups, treated as orders in PPG I, and that these, both together and individually, are monophyletic, being related as shown in the cladogram below:


Classification

The rank and name used for the taxon holding the extant lycophytes (and their closest extinct relatives) varies widely. Table 1 below shows some of the highest ranks that have been used. Systems may use taxa at a rank lower than the highest given in the table with the same circumscription; for example, a system that uses Lycopodiophyta as the highest ranked taxon may place all of its members in a single subclass. Some systems use a higher rank for a more broadly defined taxon of lycophytes that includes some extinct groups more distantly related to extant lycophytes, such as the
zosterophyll The zosterophylls are a group of extinct land plants that first appeared in the Silurian period. The taxon was first established by Banks in 1968 as the subdivision Zosterophyllophytina; they have since also been treated as the division Zoste ...
s. For example, Kenrick & Crane (1997) use the subdivision Lycophytina for this purpose, with all extant lycophytes falling within the class Lycopsida. Other sources exclude the zosterophylls from any "lycophyte" taxon. In the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relatio ...
classification of 2016 (PPG I), the three orders are placed in a single class, Lycopodiopsida, holding all extant lycophyte species. Older systems have used either three classes, one for each order, or two classes, recognizing the closer relationship between Isoetales and Selaginellales. In these cases, a higher ranked taxon is needed to contain the classes (see Table 1). As Table 2 shows, the names "Lycopodiopsida" and "Isoetopsida" are both ambiguous.


Subdivisions

The PPG I system divides up the extant lycophytes as shown below. *Class Lycopodiopsida Bartl. (3 orders) :*Order
Lycopodiales The Lycopodiaceae (class Lycopodiopsida, order Lycopodiales) are an old family of vascular plants, including all of the core clubmosses and firmosses, comprising 16 accepted genera and about 400 known species. This family originated about 380 mil ...
DC. ex Bercht. & J.Presl (1 extant family) ::*Family
Lycopodiaceae The Lycopodiaceae (class Lycopodiopsida, order Lycopodiales) are an old family of vascular plants, including all of the core clubmosses and firmosses, comprising 16 accepted genera and about 400 known species. This family originated about 380 m ...
P.Beauv. (16 extant genera) :*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 (1 extant family) ::*Family
Isoetaceae Isoetaceae is a family including living quillworts (''Isoetes'') and comparable extinct herbaceous lycopsids (''Tomiostrobus ''Tomiostrobus'' is an extinct quillwort genus from the Early Triassic of Australia, China and Russia, which was espe ...
Dumort. (1 extant genus) :*Order
Selaginellales ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
Prantl (1 extant family) ::*Family
Selaginellaceae ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
Willk (1 extant genus) Some extinct groups, such as
zosterophyll The zosterophylls are a group of extinct land plants that first appeared in the Silurian period. The taxon was first established by Banks in 1968 as the subdivision Zosterophyllophytina; they have since also been treated as the division Zoste ...
s, fall outside the limits of the taxon as defined by the classifications in Table 1 above. However, other extinct groups fall within some circumscriptions of this taxon. Taylor et al. (2009) and Mauseth (2014) include a number of extinct orders in their division (phylum) Lycophyta, although they differ on the placement of some genera. The orders included by Taylor et al. are: *Order † Drepanophycales (including '' Baragwanathia'', ''
Drepanophycus ''Drepanophycus'' is a genus of extinct plants of the division Lycopodiophyta of Early to Late Devonian age (around ), found in Eastern Canada and Northeast US, China, Russia, Egypt and various parts of Northern Europe and Britain. Description ...
'' and ''
Asteroxylon ''Asteroxylon'' ("star-shaped xylem") is an extinct genus of vascular plants of the Division Lycopodiophyta known from anatomically preserved specimens described from the famous Early Devonian Rhynie chert and Windyfield chert in Aberdeenshire, ...
'') *Order †
Protolepidodendrales The Protolepidodendrales are an extinct order of lycopsids that flourished from the Devonian to the lower Carboniferous ( Mississippian) periods. '' Leclercqia'' is one of the best-known genera. '' Protolepidodendropsis'' is another genus incl ...
*Order †
Lepidodendrales Lepidodendrales (from the Greek for "scale tree") were primitive, vascular, heterosporous, arborescent (tree-like) plants related to present day lycopsids. Members of Lepidodendrales are the best understood of the fossil lycopsids due to the vas ...
*Order † Pleuromeiales Mauseth uses the order †Asteroxylales, placing ''Baragwanathia'' in the Protolepidodendrales. The relationship between some of these extinct groups and the extant ones was investigated by Kenrick and Crane in 1997. When the genera they used are assigned to orders, their suggested relationship is:


Evolution

The Lycopodiopsida are distinguished from other vascular plants by the possession of microphylls and by their sporangia, which are lateral as opposed to terminal and which open (dehisce) transversely rather than longitudinally. In some groups, the sporangia are borne on sporophylls that are clustered into strobili. Phylogenetic analysis shows the group branching off at the base of the evolution of vascular plants and they have a long evolutionary history.
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 preserved ...
s are abundant worldwide, especially in coal deposits. Fossils that can be ascribed to the Lycopodiopsida first appear in the Silurian period, along with a number of other vascular plants. The Silurian '' Baragwanathia longifolia'' is one of the earliest identifiable species. ''
Lycopodolica ''Lycopodolica'' is a genus of extinct plants of the Late Silurian (, around ). Fossils were found in the Rashkov Beds in Podolia in modern Ukraine. Plants there are preserved as compressions without internal detail. ''Lycopodolica'' had stems ...
'' is another Silurian genus which appears to be an early member of this group. The group evolved roots independently from the rest of the vascular plants. From the Devonian onwards, some species grew large and tree-like. Devonian fossil trees from Svalbard, growing in equatorial regions, raise the possibility that they drew down enough carbon dioxide to change the earth's climate significantly. During the Carboniferous, tree-like forms (such as ''
Lepidodendron ''Lepidodendron'' is an extinct genus of primitive vascular plants belonging to the family Lepidodendraceae, part of a group of Lycopodiopsida known as scale trees or arborescent lycophytes, related to quillworts and lycopsids (club mosses). Th ...
'' and other "scale-trees" of the order
Lepidodendrales Lepidodendrales (from the Greek for "scale tree") were primitive, vascular, heterosporous, arborescent (tree-like) plants related to present day lycopsids. Members of Lepidodendrales are the best understood of the fossil lycopsids due to the vas ...
) formed huge forests that dominated the landscape. Unlike modern trees, leaves grew out of the entire surface of the trunk and branches, but fell off as the plant grew, leaving only a small cluster of leaves at the top. The trees are marked with diamond-shaped scars where they once had leaves. Quillworts (order Isoetales) and Selaginella are considered their closest extant relatives and share some unusual features with these
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 preserved ...
trees, including the development of both bark,
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from w ...
and
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...
, a modified shoot system acting as roots, bipolar and
secondary growth In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of ...
, and an upright stance. The remains of scale-trees formed many fossil
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when ...
deposits. In Fossil Park,
Glasgow Glasgow ( ; sco, Glesca or ; gd, Glaschu ) is the most populous city in Scotland and the fourth-most populous city in the United Kingdom, as well as being the 27th largest city by population in Europe. In 2020, it had an estimated popul ...
,
Scotland Scotland (, ) is a Countries of the United Kingdom, country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a Anglo-Scottish border, border with England to the southeast ...
, fossilized lycophyte trees can be found in
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates ...
. The Lycopodiopsida had their maximum diversity in the Pennsylvanian (Upper Carboniferous), particularly tree-like ''Lepidodendron'' and ''
Sigillaria ''Sigillaria'' is a genus of extinct, spore-bearing, arborescent (tree-like) plants. It was a lycopodiophyte, and is related to the lycopsids, or club-mosses, but even more closely to quillworts, as was its associate ''Lepidodendron''. Fossil ...
'' that dominated tropical wetlands. The complex ecology of these tropical rainforests collapsed during the Middle Pennsylvanian due to a change in climate. In Euramerica, tree-like species apparently became extinct in the Late Pennsylvanian, as a result of a transition to a much drier climate, giving way to conifers,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s and
horsetails ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a " living fossil", the only living genus of the entire subclass ...
. In
Cathaysia Cathaysia was a microcontinent or a group of terranes that rifted off Gondwana during the Late Paleozoic. They mostly correspond to modern territory of China, which were split into the North China and South China blocks. Terminology The terms " ...
(now South China), tree-like species survived into the
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last ...
. Nevertheless, lycopodiopsids are rare in the
Lopingian The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic. The Lopingian is often synonymous with the informal terms l ...
(latest Permian), but regained dominance in the
Induan The Induan is the first age of the Early Triassic epoch in the geologic timescale, or the lowest stage of the Lower Triassic series in chronostratigraphy. It spans the time between 251.902 Ma and Ma (million years ago). The Induan is sometime ...
(earliest Triassic), particularly '' Pleuromeia''. After the worldwide
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
, members of this group pioneered the repopulation of habitats as opportunistic plants. The heterogeneity of the terrestrial plant communities increased markedly during the Middle Triassic when plant groups like horsetails, ferns, pteridosperms,
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
s, ginkgos and conifers resurfaced and diversified quickly.


Microbial associations

Lycophytes form associations with microbes such as fungi and bacteria, including
arbuscular mycorrhiza An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscules. ( ...
l and
endophytic An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
associations. Arbuscular mycorrhizal associations have been characterized in all stages of the lycophyte lifecycle:
mycoheterotrophic Myco-heterotrophy (from Greek μύκης , "fungus", ἕτερος ', "another", "different" and τροφή ', "nutrition") is a symbiotic relationship between certain kinds of plants and fungi, in which the plant gets all or part of its food fro ...
gametophyte, photosynthetic surface-dwelling gametophyte, young sporophyte, and mature sporophyte. Arbuscular mycorrhizae have been found in ''
Selaginella ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
spp.'' roots and vesicles. During the mycoheterotrophic gametophyte lifecycle stage, lycophytes gain all of their carbon from subterranean glomalean fungi. In other plant taxa, glomalean networks transfer carbon from neighboring plants to mycoheterotrophic gametophytes. Something similar could be occurring in ''Huperzia hypogeae'' gametophytes which associate with the same glomalean phenotypes as nearby ''Huperzia hypogeae'' sporophytes. Fungal endophytes have been found in many species of lycophyte, however the function of these endophytes in host plant biology is not known. Endophytes of other plant taxa perform roles such as improving plant competitive fitness, conferring biotic and abiotic stress tolerance, promoting plant growth through phytohormone production or production of limiting nutrients. However, some endophytic fungi in lycophytes do produce medically relevant compounds. ''Shiraia'' sp Slf14 is an endophytic fungus present in ''
Huperzia serrata ''Huperzia serrata'', the toothed clubmoss, is a plant known as a firmoss which contains the acetylcholinesterase inhibitor huperzine A Huperzine A is a naturally occurring sesquiterpene alkaloid compound found in the firmoss ''Huperzia serr ...
'' that produces
Huperzine A Huperzine A is a naturally occurring sesquiterpene alkaloid compound found in the firmoss ''Huperzia serrata'' and in varying quantities in other food ''Huperzia'' species, including ''H. elmeri'', ''H. carinat'', and ''H. aqualupian''. Huperz ...
, a biomedical compound which has been approved as a drug in China and a dietary supplement in the U.S. to treat Alzheimer's Disease. This fungal endophyte can be cultivated much more easily and on a much larger scale than ''H. serrata'' itself which could increase the availability of Huperzine A as a medicine.


Uses

The spores of lycopods are highly flammable and so have been used in
fireworks Fireworks are a class of low explosive pyrotechnic devices used for aesthetic and entertainment purposes. They are most commonly used in fireworks displays (also called a fireworks show or pyrotechnics), combining a large number of devices ...
.
Lycopodium powder Lycopodium powder is a yellow-tan dust-like powder, consisting of the dry spores of clubmoss plants, or various fern relatives. When it is mixed with air, the spores are highly flammable and are used to create dust explosions as theatrical spec ...
, the dried spores of the common clubmoss, was used in Victorian theater to produce flame-effects. A blown cloud of spores burned rapidly and brightly, but with little heat. (It was considered safe by the standards of the time.)


References


External links


Introduction to the Lycophyta
from the University of California Museum of Paleontology {{Authority control Lycophytes Plant classes