''Huperzia'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

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 i ...

'', 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 (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...

(''Abies''), a

conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...

. , 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 relation ...

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 mil ...

. Most species in the subfamily are placed in the genus ''

Phlegmariurus ''Phlegmariurus'' is a genus of lycophyte plants in the family Lycopodiaceae. The genus is recognized in the Pteridophyte Phylogeny Group classification of 2016 (PPG I), but not by some other sources, which keep it in a broadly defined ''Huperzia ...

''. ''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

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 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 cy ...

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

mycorrhiza A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...

l. 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 Johann Jakob Bernhardi (1 September 1774, in Erfurt – 13 May 1850, in Erfurt) was a German doctor and botanist. Biography Johann J. Bernhardi studied Medicine and Botany at the University of Erfurt, and after graduation practiced medicine for a ...

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 ''Huperzia selago'', the northern firmoss or fir clubmoss, is a vascular plant in the family Lycopodiaceae. It has a circumpolar distribution in temperate and boreal regions in both hemispheres. Distribution and habitat It is found in the north ...

''. 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 gro ...

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

'' 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 centra ...

''. 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 ''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 ...

'' Beitel & Mickel *''

Huperzia archboldiana ''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 ...

'' (Nessel) Holub *''

Huperzia arunachalensis ''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 ...

'' (D.D.Pant & P.S.Pandey) Fraser-Jenk. *''

Huperzia asiatica ''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) N.Shrestha & X.C.Zhang *''

Huperzia australiana ''Huperzia australiana'' is a species of small terrestrial plant, a firmoss, in the Lycopodiaceae (clubmoss) family. It is native to Australia and New Zealand. Distribution and habitat The plant occurs at sheltered sites in subalpine and subant ...

'' (Herter) Holub *''

Huperzia beccarii ''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 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 ''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 ...

'' X.Y.Wang & P.S.Wang *'' Huperzia continentalis'' 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'' (Herter) Holub *'' Huperzia fuegiana'' (Roiv.) Holub *'' Huperzia gedeana'' (Alderw.) Holub *'' Huperzia goliathensis'' (Alderw.) Holub *'' Huperzia haleakalae'' (Brack.) Holub *'' Huperzia herteriana'' (Kümmerle) T.Sen & U.Sen *'' Huperzia javanica'' (Sw.) Fraser-Jenk. *'' Huperzia jejuensis'' B.Y.Sun & J.Lim *'' Huperzia kangdingensis'' (Ching) Ching *'' Huperzia kunmingensis'' Ching *'' Huperzia laipoensis'' Ching *'' Huperzia lajouensis'' Ching *'' Huperzia leishanensis'' X.Y.Wang *'' Huperzia liangshanica'' (H.S.Kung) Ching & H.S.Kung *''

Huperzia lucidula ''Huperzia lucidula'', the shining firmoss or shining clubmoss, is a bright evergreen, rhizomatous clubmoss of the genus ''Huperzia''. They grow in loose tufts 14–20 cm long, occasionally up to 1 m long. The leaves are 7–11 mm lo ...

'' (Michx.) Trevis. *'' Huperzia medogensis'' Ching & Y.X.Lin *'' Huperzia meghalaica'' Fraser-Jenk. *'' Huperzia miniata'' (Spring) Trevis. *'' Huperzia minima'' (Herter) Holub *'' Huperzia miyoshiana'' (Makino) Ching *'' Huperzia muscicola'' W.M.Chu *'' Huperzia nanchuanensis'' (Ching & H.S.Kung) Ching & H.S.Kung *'' Huperzia nanlingensis'' Shrestha, F.W.Xing, X.P.Qi, Y.H.Yan & X.C.Zhang *'' Huperzia occidentalis'' (Clute) Kartesz & Gandhi *'' Huperzia petrovii'' Sipliv. *''

Huperzia porophila ''Huperzia porophila'', the rock clubmoss or rock firmoss, grows throughout the Appalachian Mountains, Appalachian province of the Eastern United States and central Canada, from Ontario south to Georgia (U.S. state), Georgia and Alabama. It is r ...

'' (F.E.Lloyd & Underw.) Holub *'' Huperzia quasipolytrichoides'' (Hayata) Ching *'' Huperzia rubicaulis'' S.K.Wu & X.Cheng *'' Huperzia rubricaulis'' (Alderw.) Holub *'' Huperzia saururoides'' (Bory & D'Urv.) Rothm. *''

'' (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 huperzine A Huperzine A is a naturally occurring sesquiterpene alkaloid compound found in the firmoss ''Huperzia serr ...

'' (Thunb.) Trevis. *'' Huperzia shresthae'' Fraser-Jenk. *'' Huperzia somae'' (Hayata) Ching *'' Huperzia sprengeri'' (Nessel) Holub *'' Huperzia suberecta'' (Lowe) Tardieu *'' Huperzia sumatrana'' (Alderw.) Holub *'' Huperzia sutchueniana'' (Herter) Ching *'' Huperzia tibetica'' (Ching) Ching *'' Huperzia yakusimensis'' (Herter) Holub *'' Huperzia zollingeri'' (Herter) Holub

Former species

Many species formerly placed in ''Huperzia'' have been transferred to the genus ''

'', 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'' *''Huperzia crassa'' (Humb. & Bonpl. ex Willd.) Rothm. = '' Phlegmariurus crassus'' *''Huperzia cumingii'' (Nessel) Holub = '' Phlegmariurus cumingii'' *''Huperzia espinosana'' B.Øllg. = '' Phlegmariurus espinosanus'' *''Huperzia hastata'' B.Øllg. = '' Phlegmariurus hastatus'' *''Huperzia llanganatensis'' B.Øllg. = '' Phlegmariurus llanganatensis'' *''Huperzia loxensis'' B.Øllg. = '' Phlegmariurus loxensis'' *''Huperzia mannii'' (Hillebr.) Holub = '' Phlegmariurus mannii'' *''Huperzia megastachya'' (Baker) Tardieu = '' Phlegmariurus megastachyus'' *''Huperzia nutans'' (Brack.) Rothm. = '' Phlegmariurus nutans'' – (

oahu Oahu () (Hawaiian language, Hawaiian: ''Oʻahu'' ()), also known as "The Gathering place#Island of Oʻahu as The Gathering Place, Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over t ...

kauai Kauai, () anglicized as Kauai ( ), is geologically the second-oldest of the main Hawaiian Islands (after Niʻihau). With an area of 562.3 square miles (1,456.4 km2), it is the fourth-largest of these islands and the 21st largest island ...

) *''Huperzia phlegmaria'' (L.) Rothm. = '' Phlegmariurus phlegmaria'' *''Huperzia polydactyla'' B.Øllg. = '' Phlegmariurus polydactylus'' *''Huperzia scabrida'' B.Øllg. = '' Phlegmariurus scabridus'' *''Huperzia talpiphila'' B.Øllg. = '' Phlegmariurus talpiphilus'' *''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 polyg ...

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

Picture gallery
{{Taxonbar, from=Q2111150 Lycophyte genera