''Huperzia'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...

lycophyte 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 ...

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 ''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 ...

'', 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 Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family Pinaceae. They are found on mountains throughout much of North and Central America, Europe, Asia, and North Africa. The genus is most closely related to ...

(''Abies''), a

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 ex ...

. , two very different circumscriptions of the genus were in use. 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), ''Huperzia'' is one of three genera in the subfamily Huperzioideae of 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 mi ...

. 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.

Morphology

The sporophytes of this genus have unbranched shoots that are generally upright and round in cross section. Horizontal stems are absent. The leaves are not borne in distinct ranks, and are usually somewhat lanceolate in shape. In some species, they vary in size according to the season in which they grow. Branchlets bearing gemmae – bud-like structures by which the plant reproduces asexually – occur among the leaves. The gemmae are triangular, with eight leaves in a constant pattern: four flattened into a plane and two large lateral leaves. The

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 ...

are kidney-shaped (reniform), occurring at the base of a leaf that is either unmodified or reduced. The roots are produced near the apex of shoots, and migrate downwards inside the cortex of the stem to emerge at soil level. The unbranched

gametophyte A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...

s are not photosynthetic, but rather subterranean and mycorrhizal. The ''Flora of North America'' distinguishes ''Huperzia'' from the

epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...

tropical genus ''Phlegmariurus'' on the basis of differences such as the former's complex and specialized shoots, the gemmae and the branchlets on which they are borne, and the unbranched gametophytes.

Taxonomy

The genus ''Huperzia'' was created by Johann Jakob Bernhardi in 1801. Bernhardi separated ''Huperzia'' from ''Lycopodium''. The

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...

is ''Lycopodium selago'' which became '' Huperzia selago''. In the

classification of 2016 (PPG I), ''Huperzia'' is placed in the subfamily Huperzioideae of the family

. A

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

study in 2016, employing both molecular and morphological data, concluded that either a one-genus or a three-genus division of the subfamily produced

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...

taxa. The authors preferred the three-genus division, recognizing ''Huperzia'', '' Phlegmariurus'' and ''

Phylloglossum ''Phylloglossum'', a genus in the clubmoss family Lycopodiaceae, is a small plant superficially resembling a tiny grass plant, growing with a rosette of slender leaves 2–5 cm long from an underground bulb-like root. It has a single centr ...

''. Their preferred hypothesis for the relationships of the three genera was: The majority of the species formerly placed in a broadly defined ''Huperzia'' belong in ''Phlegmariurus''. Earlier, the ''Flora of North America'' had also separated ''Huperzia'' from ''Phegmariurus''. However, ''Phlegmariurus'' is difficult to separate morphologically, and others have preferred the one-genus division of the subfamily.

Species

The PPG I classification stated there were 25 species in the genus ''Huperzia''. , ''World Ferns'' listed 56 species, noting that "many species still need transfer into other split genera". *'' Huperzia appalachiana'' Beitel & Mickel *'' Huperzia archboldiana'' (Nessel) Holub *'' Huperzia arunachalensis'' (D.D.Pant & P.S.Pandey) Fraser-Jenk. *'' Huperzia asiatica'' (Ching) N.Shrestha & X.C.Zhang *'' Huperzia australiana'' (Herter) Holub *'' Huperzia beccarii'' (Alderw.) Holub *''

Huperzia bucahwangensis ''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 dif ...

'' Ching *''

Huperzia campestris ''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 dif ...

'' (Alderw.) Holub *''

Huperzia chinensis ''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 dif ...

'' (Christ) Ching *'' Huperzia chishuiensis'' X.Y.Wang & P.S.Wang *''

Huperzia continentalis ''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 d ...

'' Testo, A.Haines & A.V.Gilman *'' Huperzia crispata'' (Ching) Ching *'' Huperzia emeiensis'' (Ching & H.S.Kung) Ching & H.S.Kung *'' Huperzia erubescens'' (Brack.) Holub *''

Huperzia everettii ''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 d ...

'' (Herter) Holub *''

Huperzia fuegiana ''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 d ...

'' (Roiv.) Holub *'' Huperzia gedeana'' (Alderw.) Holub *'' Huperzia goliathensis'' (Alderw.) Holub *''

Huperzia haleakalae ''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 d ...

'' (Brack.) Holub *''

Huperzia herteriana ''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 d ...

'' (Kümmerle) T.Sen & U.Sen *''

Huperzia javanica ''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 ...

'' (Sw.) Fraser-Jenk. *''

Huperzia jejuensis ''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 ...

'' B.Y.Sun & J.Lim *''

Huperzia kangdingensis ''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 ...

'' (Ching) Ching *''

Huperzia kunmingensis ''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 d ...

'' Ching *''

Huperzia laipoensis ''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 d ...

'' Ching *''

Huperzia lajouensis ''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 d ...

'' Ching *''

Huperzia leishanensis ''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 d ...

'' X.Y.Wang *'' Huperzia liangshanica'' (H.S.Kung) Ching & H.S.Kung *'' Huperzia lucidula'' (Michx.) Trevis. *'' Huperzia medogensis'' Ching & Y.X.Lin *''

Huperzia meghalaica ''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 d ...

'' Fraser-Jenk. *''

Huperzia miniata ''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 ...

'' (Spring) Trevis. *'' Huperzia minima'' (Herter) Holub *'' Huperzia miyoshiana'' (Makino) Ching *''

Huperzia muscicola ''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 d ...

'' W.M.Chu *'' Huperzia nanchuanensis'' (Ching & H.S.Kung) Ching & H.S.Kung *''

Huperzia nanlingensis ''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 d ...

'' Shrestha, F.W.Xing, X.P.Qi, Y.H.Yan & X.C.Zhang *''

Huperzia occidentalis ''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 d ...

'' (Clute) Kartesz & Gandhi *'' Huperzia petrovii'' Sipliv. *'' Huperzia porophila'' (F.E.Lloyd & Underw.) Holub *'' Huperzia quasipolytrichoides'' (Hayata) Ching *''

Huperzia rubicaulis ''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 d ...

'' S.K.Wu & X.Cheng *'' Huperzia rubricaulis'' (Alderw.) Holub *'' Huperzia saururoides'' (Bory & D'Urv.) Rothm. *'' Huperzia selago'' (L.) Bernh. (including ''H. selago'' subsp. ''appressa'' - Appalachian firmoss) *''

Huperzia serrata ''Huperzia serrata'', the toothed clubmoss, is a plant known as a firmoss which contains the acetylcholinesterase inhibitor Inhibitor or inhibition may refer to: In biology * Enzyme inhibitor, a substance that binds to an enzyme and decreases ...

'' (Thunb.) Trevis. *'' Huperzia shresthae'' Fraser-Jenk. *'' Huperzia somae'' (Hayata) Ching *'' Huperzia sprengeri'' (Nessel) Holub *''

Huperzia suberecta ''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 ...

'' (Lowe) Tardieu *''

Huperzia sumatrana ''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 ...

'' (Alderw.) Holub *'' Huperzia sutchueniana'' (Herter) Ching *''

Huperzia tibetica ''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 ...

'' (Ching) Ching *''

Huperzia yakusimensis ''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 ...

'' (Herter) Holub *'' Huperzia zollingeri'' (Herter) Holub

Former species

Many species formerly placed in ''Huperzia'' have been transferred to the genus '' Phlegmariurus'', including the following (synonyms from ''World Ferns''): *''Huperzia ascendens'' = '' Phlegmariurus ascendens'' *''Huperzia austroecuadorica'' B.Øllg. = '' Phlegmariurus austroecuadoricus'' *''Huperzia columnaris'' B.Øllg. = '' Phlegmariurus columnaris'' *''Huperzia compacta'' (Hook.) Trevis = ''

Phlegmariurus compactus ''Phlegmariurus compactus'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador, where it is distributed throughout the southern Andean páramo Páramo () can refer to a variety of alpine tundra ecosystems located in t ...

'' *''Huperzia crassa'' (Humb. & Bonpl. ex Willd.) Rothm. = ''

Phlegmariurus crassus ''Phlegmariurus crassus'' is a lycophyte found in Central and South America. Distribution It is found from Hispaniola and Mexico south through South America to Paraguay. References crassus Marcus Licinius Crassus (; 115 – 53 BC) was ...

'' *''Huperzia cumingii'' (Nessel) Holub = ''

Phlegmariurus cumingii ''Phlegmariurus cumingii'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador. Its natural habitats are subtropical or tropical moist montane forest and subtropical or tropical high-altitude grassland A grassland i ...

'' *''Huperzia espinosana'' B.Øllg. = ''

Phlegmariurus espinosanus ''Phlegmariurus espinosanus'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador. Its natural habitat is subtropical or tropical high-altitude grassland. It is threatened by habitat loss Habitat destruction (also t ...

'' *''Huperzia hastata'' B.Øllg. = ''

Phlegmariurus hastatus ''Phlegmariurus hastatus'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador, where it is known from two locations in Loja Province. It grows on the páramo Páramo () can refer to a variety of alpine tundra ecosy ...

'' *''Huperzia llanganatensis'' B.Øllg. = '' Phlegmariurus llanganatensis'' *''Huperzia loxensis'' B.Øllg. = ''

Phlegmariurus loxensis ''Phlegmariurus loxensis'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador, where it occurs in the páramo Páramo () can refer to a variety of alpine tundra ecosystems located in the Andes Mountain Range, South ...

'' *''Huperzia mannii'' (Hillebr.) Holub = ''

Phlegmariurus mannii ''Phlegmariurus mannii'', synonym ''Huperzia mannii'', is a species of lycopod, known by the common names Mann's clubmoss and ''wawaeiole''. It is endemic to Hawaii, where there are only six populations remaining. It is a federally listed endange ...

'' *''Huperzia megastachya'' (Baker) Tardieu = '' Phlegmariurus megastachyus'' *''Huperzia nutans'' (Brack.) Rothm. = ''

Phlegmariurus nutans ''Phlegmariurus nutans'', synonym ''Huperzia nutans'', known as wawaeiole or nodding clubmoss, is a species of club moss in the family Lycopodiaceae. It is endemic to wet forests and cliffside shrublands on the islands of Oahu and Kauai ...

'' – (

oahu Oahu () ( Hawaiian: ''Oʻahu'' ()), also known as "The Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over two-thirds of the population of the U.S. state of Hawaii. The island of O� ...

, kauai) *''Huperzia phlegmaria'' (L.) Rothm. = '' Phlegmariurus phlegmaria'' *''Huperzia polydactyla'' B.Øllg. = ''

Phlegmariurus polydactylus ''Phlegmariurus polydactylus'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador. Its natural habitats are subtropical or tropical moist montane forest and subtropical or tropical high-altitude grassland. It is threaten ...

'' *''Huperzia scabrida'' B.Øllg. = ''

Phlegmariurus scabridus ''Phlegmariurus scabridus'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador. Its natural habitats are subtropical or tropical moist montane forest and subtropical or tropical high-altitude grassland. It is threatened ...

'' *''Huperzia talpiphila'' B.Øllg. = ''

Phlegmariurus talpiphilus ''Phlegmariurus talpiphilus'' is a species of plant in the family Lycopodiaceae. It is endemic to Ecuador. Its natural habitat is subtropical or tropical high-altitude grassland. It is threatened by habitat loss Habitat destruction (also t ...

'' *''Huperzia varia'' (R.Br.) Trevis. = '' Phlegmariurus varius''

Distribution and habitat

circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every po ...

in the PPG I classification, ''Huperzia'' is distributed in temperate, arctic and alpine habitats, including mountains in tropical Asia. Its species are terrestrial or grow on rocks. ''Phlegmariurus'' is

, and has a worldwide tropical distribution, so when ''Huperzia'' is defined broadly to include all three genera of the subfamily Huperzioideae, it has an almost worldwide distribution, absent mainly in North Africa, the Arabian Peninsula and Western Asia.

References

External links

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{{Taxonbar, from=Q2111150 Lycophyte genera