Lycopodiopsida
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Lycopodiopsida is a class of vascular plants 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 and reproduce by means of
spore In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells tha ...
s borne in sporangia on the sides of the stems at the bases of the leaves. Although living species are small, during the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonif ...
, 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 stratum, rock strata called coal seams. Coal is mostly carbon with variable amounts of other Chemical element, elements, chiefly hydrogen, sulfur, oxygen, and nitrogen ...
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 (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, Isoetales and Selaginellales.


Characteristics

Club-mosses (Lycopodiales) are homosporous, but the genera '' Selaginella'' (spikemosses) and '' Isoetes'' (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 Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduct ...
(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; 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 gametophytes are mycoheterotrophic and long-lived, residing underground for several years before emerging from the ground and progressing to the
sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant Plants are predominantly photosynthetic eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many un ...
stage.


Taxonomy


Phylogeny

The extant lycophytes are vascular plants (tracheophytes) with microphyllous leaves, distinguishing them from the euphyllophytes (plants with megaphyllous leaves). The sister group of the extant lycophytes and their closest extinct relatives are generally believed to be the zosterophylls, a paraphyletic or plesion 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 In cladistics for a group of organisms, monophyly is the condition of being a clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common anc ...
, 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 zosterophylls. 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 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 DC. ex Bercht. & J.Presl (1 extant family) ::*Family Lycopodiaceae P.Beauv. (16 extant genera) :*Order Isoetales Prantl (1 extant family) ::*Family Isoetaceae Dumort. (1 extant genus) :*Order Selaginellales Prantl (1 extant family) ::*Family Selaginellaceae Willk (1 extant genus) Some extinct groups, such as zosterophylls, 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'' and '' Asteroxylon'') *Order † Protolepidodendrales *Order † Lepidodendrales *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 Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire The Roman Empire ( la, Imperium Romanum ; grc-gre, ...
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 The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian The Devonian ( ) is a period (geology), geologic period an ...
'' Baragwanathia longifolia'' is one of the earliest identifiable species. '' Lycopodolica'' 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 The Devonian ( ) is a period (geology), geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Dev ...
onwards, some species grew large and tree-like. Devonian fossil trees from
Svalbard Svalbard ( , ), also known as Spitsbergen, or Spitzbergen, is a Norway, Norwegian archipelago in the Arctic Ocean. North of continental Europe, mainland Europe, it is about midway between the northern coast of Norway and the North Pole. The ...
, growing in equatorial regions, raise the possibility that they drew down enough carbon dioxide to change the earth's climate significantly. During the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonif ...
, tree-like forms (such as '' Lepidodendron'' and other "scale-trees" of the order Lepidodendrales) 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 Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire The Roman Empire ( la, Imperium Romanum ; grc-gre, ...
trees, including the development of both bark, cambium and wood, a modified shoot system acting as roots, bipolar and secondary growth, 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 stratum, rock strata called coal seams. Coal is mostly carbon with variable amounts of other Chemical element, elements, chiefly hydrogen, sulfur, oxygen, and nitrogen ...
deposits. In Fossil Park,
Glasgow Glasgow ( ; sco, Glesca or ; gd, Glaschu ) is the most populous city in Scotland Scotland (, ) is a country that is part of the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as ...
,
Scotland Scotland (, ) is a country that is part of the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the ...
, 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 rock Sedimentary rocks are types of rock that are formed by the accu ...
. The Lycopodiopsida had their maximum diversity in the Pennsylvanian (Upper Carboniferous), particularly tree-like ''Lepidodendron'' and '' Sigillaria'' that dominated tropical wetlands. The complex ecology of these tropical rainforests collapsed during the Middle Pennsylvanian due to a change in climate. In
Euramerica Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana Gondwana () was a large landmass A landmass, or land mass, is a large region or ...
, tree-like species apparently became extinct in the Late Pennsylvanian, as a result of a transition to a much drier climate, giving way to
conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
s,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spore In biology Biology is the scientific study of life. It is a natural science wit ...
s and horsetails. In Cathaysia (now South China), tree-like species survived into the Permian. Nevertheless, lycopodiopsids are rare in the Lopingian (latest Permian), but regained dominance in the Induan (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 The Triassic ( ) ...
, 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 ma ...
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 A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes m ...
l and endophytic associations. Arbuscular mycorrhizal associations have been characterized in all stages of the lycophyte lifecycle: mycoheterotrophic gametophyte, photosynthetic surface-dwelling gametophyte, young sporophyte, and mature sporophyte. Arbuscular mycorrhizae have been found in '' Selaginella 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'' that produces Huperzine A, 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. Lycopodium powder, 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