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''Lycopodioideae'' is a subfamily in the 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 mil ...
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 relation ...
classification of 2016 (PPG I). It is equivalent to a broad
circumscription Circumscription may refer to: *Circumscribed circle *Circumscription (logic) *Circumscription (taxonomy) * Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...
of the genus ''
Lycopodium ''Lycopodium'' (from Greek ''lykos'', wolf and ''podion'', diminutive of ''pous'', foot) is a genus of clubmosses, also known as ground pines or creeping cedars, in the family Lycopodiaceae. Two very different circumscriptions of the genus are i ...
'' in other classifications. Like all
lycophytes The lycophytes, when broadly circumscribed, are a vascular plant (tracheophyte) subgroup of the kingdom Plantae. They are sometimes placed in a division Lycopodiophyta or Lycophyta or in a subdivision Lycopodiophytina. They are one of the oldest ...
, members of the Lycopodioideae reproduce by spores. The oldest fossils of modern members of the family date to the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
.


Description

The
sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote pr ...
s of Lycopodioideae species are relatively short herbaceous plants. They have stems with pseudomonopodial branching in which unequal binary branching produces the appearance of a main stem with secondary side branches. The main stems are indeterminate and of various forms, including
rhizomatous In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
, creeping, trailing and climbing. They usually form roots at intervals along their length. The branches are usually determinate (i.e. of limited growth and extension).
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 cy ...
are borne at the bases or in the axils of special spore-bearing leaves (
sporophyll A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls (whether they are microphylls or megaphylls) bear either megasporangia and thus are called megasporophylls, or mi ...
s), which are notably different from the normal leaves, and are usually grouped into compact terminal structures (
strobili A strobilus (plural: strobili) is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to th ...
). The strobili may be either upright or drooping.


Taxonomy

The family Lycopodiaceae was first established in 1802. Although other genera now placed within the family (in particular ''
Huperzia ''Huperzia'' is a genus of Lycopodiopsida, lycophyte plants, sometimes known as the firmosses or fir clubmosses; the ''Flora of North America'' calls them gemma fir-mosses. This genus was originally included in the related genus ''Lycopodium'', f ...
'', published in 1801) had been described, until the mid-1900s, ''
Lycopodium ''Lycopodium'' (from Greek ''lykos'', wolf and ''podion'', diminutive of ''pous'', foot) is a genus of clubmosses, also known as ground pines or creeping cedars, in the family Lycopodiaceae. Two very different circumscriptions of the genus are i ...
'' was often the only genus recognized. Work by
Josef Holub Josef may refer to *Josef (given name) *Josef (surname) Josef is the surname of the following people: * Jens Josef (born 1967), German composer of classical music, a flutist and academic teacher * Michelle Josef (born 1954), Canadian musician and tr ...
and
Benjamin Øllgaard Benjamin ( he, ''Bīnyāmīn''; "Son of (the) right") blue letter bible: https://www.blueletterbible.org/lexicon/h3225/kjv/wlc/0-1/ H3225 - yāmîn - Strong's Hebrew Lexicon (kjv) was the last of the two sons of Jacob and Rachel (Jacob's thir ...
in the 1980s established three clear divisions within the family. This has since been supported by
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 ...
studies. Several different ways of representing this situation taxonomically have been used, and are still in use , including three subfamilies with multiple genera, and three genera with multiple subgeneric divisions. Three subfamilies, including Lycopodioideae, were first suggested by Warren Wagner Jr. and Joseph Beitel in 1992, but were not validly published under the ''
International Code of Botanical Nomenclature The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) 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 those "trad ...
'' as it was then. The names were validated by Benjamin Øllgaard in 2015. The entire subfamily Lycopodioideae 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 relation ...
classification of 2016 (PPG I) corresponds to the single genus ''Lycopodium'' in other classifications.


Phylogeny

Within the family Lycopodiaceae, there is support for three subgroups. In 2016, Field et al. proposed that the primary division is between Lycopodielloideae plus Lycopodioideae (which comprised their Lycopodioideae) and Huperzioideae (subfamilies ''sensu'' PPG I). Field et al. (2016) included 22 species of Lycopodioideae in their analysis, which suggested the relationships among the genera shown in the following cladogram, where the number of species included in the study is shown in parentheses:


Genera

In the Pteridophyte Phylogeny Group classification of 2016 (PPG I) the Lycopodioideae has the following genera: *''
Austrolycopodium ''Austrolycopodium'' is a genus of lycophytes in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into ...
'' Holub *''
Dendrolycopodium The genus ''Dendrolycopodium'' is a clubmoss genus in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it ...
'' A.Haines *''
Diphasiastrum ''Diphasiastrum'' is a genus of clubmosses in the plant family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is placed in the subfamily Lycopodioideae. It is closely related to the genus '' Lycopodium'' ...
'' Holub *''
Diphasium ''Diphasium'' is a genus of lycophytes in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into ''Lyc ...
'' C.Presl ex Rothm. *'' Lycopodiastrum'' Holub ex R.D.Dixit *''
Lycopodium ''Lycopodium'' (from Greek ''lykos'', wolf and ''podion'', diminutive of ''pous'', foot) is a genus of clubmosses, also known as ground pines or creeping cedars, in the family Lycopodiaceae. Two very different circumscriptions of the genus are i ...
'' L. *'' Pseudodiphasium'' Holub *''
Pseudolycopodium ''Pseudolycopodium'' is a genus of lycophyte in the family Lycopodiaceae with only one species, ''Pseudolycopodium densum'', known as the bushy clubmoss. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the genus is placed in ...
'' Holub *'' Spinulum'' A.Haines All of these genera are submerged into a single genus ''Lycopodium''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
in other systems of classification. Lycopodium magellanicum - panoramio.jpg, ''
Austrolycopodium magellanicum ''Austrolycopodium magellanicum'', Synonym (taxonomy), synonym ''Lycopodium magellanicum'', the Magellanic clubmoss, is a species of vascular plant in the club moss family Lycopodiaceae. The genus ''Austrolycopodium'' is accepted in the Pteridoph ...
'' Lycopodium hickeyi.JPG, ''
Dendrolycopodium hickeyi ''Dendrolycopodium hickeyi'' (Synonym (taxonomy), synonym ''Lycopodium hickeyi'') known as Hickey's tree club-moss or Pennsylvania clubmoss, is a North American species of clubmoss in the family Lycopodiaceae. It is native to eastern and Central ...
'' Diphasiastrum alpinum.jpg, ''
Diphasiastrum alpinum ''Diphasiastrum alpinum'', the alpine clubmoss, is a species of clubmoss. It was first described by Carl Linnaeus in his Flora Lapponica, 1737, from specimens obtained in Finland. Distribution It has a circumpolar distribution across much of the ...
'' Diphasium scariosum kz1.jpg, ''
Diphasium scariosum ''Diphasium scariosum'', synonym ''Lycopodium scariosum'', commonly known as spreading clubmoss or creeping club moss, is a species in the club moss family Lycopodiaceae. The genus '' Diphasium'' is accepted in the Pteridophyte Phylogeny Group c ...
'' Lycopodium_clavatum_151207.jpg, ''
Lycopodium clavatum ''Lycopodium clavatum'' (common club moss, stag's-horn clubmoss, running clubmoss, or ground pineBailey, L.H.; Bailey, E.Z.; the staff of the Liberty Hyde Bailey Hortorium. 1976. ''Hortus third: A concise dictionary of plants cultivated in the Un ...
'' Lycopodium deuterodensum Gloucester Tops.jpg, '' Pseudolycopodium densum'' Lycopodium annotinum 161102a.jpg, ''
Spinulum annotinum ''Spinulum annotinum'', synonym ''Lycopodium annotinum'', known as interrupted club-moss, or stiff clubmoss, is a species of clubmoss native to forests of the colder parts of North America (Greenland, St. Pierre & Miquelon, all 10 provinces an ...
''


References

{{Taxonbar, from1=Q30109303 Lycopodiaceae Plant subfamilies