Lycopodioideae

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Lycopodioideae
Temporal range: Early Cretaceous–Recent
Lycopodium dendroideum.JPG
Dendrolycopodium dendroideum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Lycophytes
Class: Lycopodiopsida
Order: Lycopodiales
Family: Lycopodiaceae
Subfamily: Lycopodioideae
W.H.Wagner & Beitel ex B.Øllg. [1]
Genera

See text.

Synonyms [2]
  • LycopodiumL. sensu Øllgaard (1987)

Lycopodioideae is a subfamily in the family Lycopodiaceae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). [1] It is equivalent to a broad circumscription of the genus Lycopodium in other classifications. Like all lycophytes, members of the Lycopodioideae reproduce by spores. The oldest fossils of modern members of the subfamily date to the Early Cretaceous. [3]

Contents

Description

The sporophytes 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, 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 are borne at the bases or in the axils of special spore-bearing leaves (sporophylls), which are notably different from the normal leaves, and are usually grouped into compact terminal structures (strobili). The strobili may be either upright or drooping. [2]

Taxonomy

The family Lycopodiaceae was first established in 1802. [4] Although other genera now placed within the family (in particular Huperzia , published in 1801 [5] ) had been described, until the mid-1900s, Lycopodium was often the only genus recognized. Work by Josef Holub and Benjamin Øllgaard in the 1980s established three clear divisions within the family. [6] This has since been supported by molecular phylogenetic studies. [7] Several different ways of representing this situation taxonomically have been used, [6] and are still in use as of 2019, 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 as it was then. The names were validated by Benjamin Øllgaard in 2015. [2] The entire subfamily Lycopodioideae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I) corresponds to the single genus Lycopodium in other classifications. [1] [8] [9]

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). [7]

Lycopodiaceae

Lycopodielloideae (Lycopodiella s.l.) 

Lycopodioideae (Lycopodium s.l.) 

Lycopodioideae sensu Field et al. (2016)

Huperzioideae (Huperzia s.l.)

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: [7]

Lycopodioideae

Lycopodiastrum (only sp.)

Pseudolycopodium (only sp.)

Pseudodiphasium (only sp.)

Austrolycopodium (2 spp.)

Dendrolycopodium (2 spp.)

Diphasium (3 spp.)

Diphasiastrum (6 spp.)

Spinulum (1 sp.)

Lycopodium (5 spp.)

Genera

In the Pteridophyte Phylogeny Group classification of 2016 (PPG I) the Lycopodioideae has the following genera: [1]

All of these genera are submerged into a single genus Lycopodium sensu lato in other systems of classification. [9]

Related Research Articles

<i>Huperzia</i> Genus of vascular plants

Huperzia is a genus of 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, from which it differs in having undifferentiated sporangial leaves, and the sporangia not formed into apical cones. The common name firmoss, used for some of the north temperate species, refers to their superficial resemblance to branches of fir (Abies), a conifer. As of 2020, two very different circumscriptions of the genus were in use. In the Pteridophyte Phylogeny Group classification of 2016, Huperzia is one of three genera in the subfamily Huperzioideae of the family Lycopodiaceae. Most species in the subfamily are placed in the genus Phlegmariurus. Huperzia is left with about 25 species, although not all have been formally transferred to other genera. Other sources recognize only Huperzia, which then has about 340 species.

<span class="mw-page-title-main">Lycopodiaceae</span> Family of vascular plants

The Lycopodiaceae 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 million years ago in the early Devonian, though the diversity within the family has been much more recent. "Wolf foot" is another common name for this family due to the resemblance of either the roots or branch tips to a wolf's paw.

<i>Lycopodium</i> Genus of vascular plants in the family Lycopodiaceae

Lycopodium 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 in use. In the Pteridophyte Phylogeny Group classification of 2016, Lycopodium is one of nine genera in the subfamily Lycopodioideae, and has from nine to 15 species. In other classifications, the genus is equivalent to the whole of the subfamily, since it includes all of the other genera. More than 40 species are accepted.

<i>Diphasiastrum</i> Genus of vascular plants in the clubmoss family Lycopodiaceae

Diphasiastrum is a genus of clubmosses in the plant family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the subfamily Lycopodioideae. It is closely related to the genus Lycopodium, and some botanists treat it within a broad view of that genus as a section, Lycopodium sect. Complanata. Some species superficially resemble diminutive gymnosperms and have been given common names such as ground-pine or ground-cedar.

<i>Lycopodiella</i> Genus of spore-bearing plants

Lycopodiella is a genus in the clubmoss family Lycopodiaceae. The genus members are commonly called bog clubmosses, describing their wetland habitat. The genus has a cosmopolitan distribution, with centers of diversity in the tropical New World and New Guinea. In the past, the genus was often incorporated within the related genus Lycopodium, but was segregated in 1964. In the Pteridophyte Phylogeny Group classification of 2016, Lycopodiella is placed in the subfamily Lycopodielloideae, along with three other genera. In this circumscription, the genus has about 15 species. Other sources use a wider circumscription, in which the genus is equivalent to the Lycopodielloideae of PPG I, in which case about 40 species and hybrids are accepted.

<span class="mw-page-title-main">Microsoroideae</span> Subfamily of ferns

Microsoroideae is a subfamily in the fern family Polypodiaceae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The subfamily is also treated as the tribe Microsoreae within a very broadly defined family Polypodiaceae sensu lato. In either treatment, it includes the previously separated tribe Lepisoreae.

<i>Dendrolycopodium</i> Genus of spore-bearing plants

The genus Dendrolycopodium is a clubmoss genus in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium. It is treated as section Obscura when retained within Lycopodium. The genus includes a discrete group of plants with similar morphologies. All have erect to semi-erect, branched stems.

<i>Austrolycopodium magellanicum</i> Species of clubmoss in the family Lycopodiaceae

Austrolycopodium magellanicum, 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 Pteridophyte Phylogeny Group classification of 2016, but not in other classifications which submerge the genus in Lycopodium.

<i>Diphasiastrum sitchense</i> Species of spore-bearing plant

Diphasiastrum sitchense, the Sitka clubmoss, is a pteridophyte species native to northern North America and northeastern Asia. It is a terrestrial herb spreading by stolons running on the surface or the ground or just slightly below the surface. Leaves are appressed, broadly lanceolate, up to 3.2 mm long. Strobili are solitary on the ends of shoots. It is known from every province in Canada, plus the US States of Alaska, Oregon, Washington, Idaho, Montana, Maine, New Hampshire, Vermont, and New York. It is also found in Greenland, St. Pierre and Miquelon, Yukon, Japan, and the Kamchatka Peninsula of Asiatic Russia. It can be found in alpine meadows, open rocky barrens, and coniferous woodlands.

<i>Pseudolycopodiella</i> Genus of spore-bearing plants

Pseudolycopodiella is a genus of non-seed plants in the Lycopodiaceae, long considered part of Lycopodium, but now recognized as a separate genus. It has 10–14 recognized species, only one in North America: Pseudolycopodiella caroliniana.

<i>Spinulum</i> Genus of spore-bearing plants

Spinulum is a genus of club mosses in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium. Spinulum annotinum is widespread in the temperate Northern Hemisphere.

Austrolycopodium erectum is a species in the club moss family Lycopodiaceae. The genus Austrolycopodium is accepted in the Pteridophyte Phylogeny Group classification of 2016, but not in other classifications which submerge the genus in Lycopodium. The species is native to Bolivia, Chile and Argentina.

<i>Palhinhaea</i> Genus of plants

Palhinhaea is a genus of lycophytes in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the subfamily Lycopodielloideae. Some sources do not recognize the genus, sinking it into Lycopodiella. Palhinhaea species are widespread in the tropics and subtropics.

<i>Austrolycopodium</i> Genus of spore-bearing plants

Austrolycopodium is a genus of lycophytes in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium. Austrolycopodium species are mostly native to the temperate southern hemisphere.

<i>Diphasium</i> Genus of spore-bearing plants

Diphasium is a genus of lycophytes in the family Lycopodiaceae. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium, others include it in Diphasiastrum. Diphasium species are mostly native to the temperate southern hemisphere, but extend northwards into Central America and the Caribbean.

Lycopodiastrum is a genus of lycophyte in the family Lycopodiaceae with only one species, Lycopodiastrum casuarinoides. In the Pteridophyte Phylogeny Group classification of 2016, the genus is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium. Lycopodiastrum casuarinoides is native to south-eastern Asia, from Tibet through China to Japan in the north, and from Sumatra to Sulawesi in the south.

<i>Pseudodiphasium</i> Genus of spore-bearing plants

Pseudodiphasium is a genus of lycophyte in the family Lycopodiaceae with only one species, Pseudodiphasium volubile. In the Pteridophyte Phylogeny Group classification of 2016, the genus is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium. Pseudodiphasium volubile is native from Peninsular Malaysia to Queensland, Australia, and has been introduced into Ecuador.

<i>Pseudolycopodium</i> Genus of spore-bearing plants

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, the genus is placed in the subfamily Lycopodioideae. Some sources do not recognize the genus, sinking it into Lycopodium. Pseudolycopodium densum is native to Australia, the North Island of New Zealand and New Caledonia. It is a spore-bearing vascular plant and grows up to a metre high. It is found in a wide variety of situations, often in high rainfall areas on sandy soils.

<span class="mw-page-title-main">Lycopodielloideae</span> Subfamily of plants

Lycopodielloideae is a subfamily in the family Lycopodiaceae in the Pteridophyte Phylogeny Group classification of 2016. It is equivalent to a broad circumscription of the genus Lycopodiella in other classifications. Like all lycophytes, members of the Lycopodielloideae are vascular plants that reproduce by spores.

<i>Lemmaphyllum</i> Genus of ferns

Lemmaphyllum is a genus of ferns in the family Polypodiaceae, subfamily Microsoroideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I).

References

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  2. 1 2 3 Øllgaard, B. (2015), "Six new species and some nomenclatural changes in neotropical Lycopodiaceae", Nordic Journal of Botany, 33 (2): 186–196, doi:10.1111/njb.00652
  3. Herrera, Fabiany; Testo, Weston L.; Field, Ashley R.; Clark, Elizabeth G.; Herendeen, Patrick S.; Crane, Peter R.; Shi, Gongle (March 2022). "A permineralized Early Cretaceous lycopsid from China and the evolution of crown clubmosses". New Phytologist. 233 (5): 2310–2322. doi: 10.1111/nph.17874 . ISSN   0028-646X. PMID   34981832. S2CID   245670357.
  4. "Lycopodiaceae P.Beauv. ex Mirb.", Plants of the World Online, Royal Botanic Gardens, Kew, retrieved 2019-12-10
  5. "Huperzia Bernh.", The International Plant Names Index , retrieved 2019-12-10
  6. 1 2 Wagner Jr., Warren H. & Beitel, Joseph M. (1992), "Generic Classification of Modern North American Lycopodiaceae", Annals of the Missouri Botanical Garden, 79 (3): 676–686, doi:10.2307/2399759, JSTOR   2399759
  7. 1 2 3 Field, Ashley R.; Testo, Weston; Bostock, Peter D.; Holtum, Joseph A.M. & Waycott, Michelle (January 2016), "Molecular Phylogenetics and the Morphology of the Lycopodiaceae Subfamily Huperzioideae Supports Three Genera: Huperzia, Phlegmariurus and Phylloglossum", Molecular Phylogenetics and Evolution, 94, Part B (Pt B): 635–57, doi:10.1016/j.ympev.2015.09.024, PMID   26493224
  8. Christenhusz, Maarten J.M. & Chase, Mark W. (2014), "Trends and concepts in fern classification", Annals of Botany, 113 (9): 571–594, doi:10.1093/aob/mct299, PMC   3936591 , PMID   24532607
  9. 1 2 "Lycopodium L.", Plants of the World Online, Royal Botanic Gardens, Kew, retrieved 2019-12-10