''Huperzia'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

lycophyte The lycophytes, when broadly circumscribed, are a group of vascular plants that include the clubmosses. They are sometimes placed in a division Lycopodiophyta or Lycophyta or in a subdivision Lycopodiophytina. They are one of the oldest lineag ...

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 Ancient 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 gen ...

'', 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 are evergreen coniferous trees belonging to the genus ''Abies'' () in the family Pinaceae. There are approximately 48–65 extant species, found on mountains throughout much of North and Central America, Eurasia, and North Africa. The genu ...

(''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 sin ...

. , two very different circumscriptions of the genus were in use. In the

Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...

classification of 2016 (PPG I), ''Huperzia'' is one of three genera in the subfamily

Huperzioideae Huperzioideae is a subfamily of lycopsids in the family Lycopodiaceae. It has sometimes been recognized as a separate family, Huperziaceae. The Pteridophyte Phylogeny Group classification of 2016 (PPG I) recognizes three extant genera: * ''Huper ...

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 17 accepted genera and about 500 known species. This family originated about 380 mi ...

. 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 one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...

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, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...

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 alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...

s are not photosynthetic, but rather subterranean and

mycorrhiza A mycorrhiza (; , 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's rhizosphere, the plant root system and its surroundings. Mycorrhizae play ...

l. The ''Flora of North America'' distinguishes ''Huperzia'' from the

epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another 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 epiphyt ...

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 International_Code_of_Zoological_Nomenclature, 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 spe ...

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

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

of the family

. A

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

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

'' and ''

Phylloglossum ''Phylloglossum'', a genus in the clubmoss Family (biology), family Lycopodiaceae, is a small plant superficially resembling a tiny Poaceae, grass plant, growing with a rosette of slender leaf, leaves 2–5 cm long from an underground bulb-l ...

''. 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 the following species, noting that "many species still need transfer into other split genera". *''

Huperzia acicularis ''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 ...

'' Björk, provisionally accepted *''

Huperzia appalachiana ''Huperzia appalachiana'' is a species of clubmoss. Description The original description is:Stems tufted to shortly decumbent (), erect portions of stem tall. Stems appear to live for definite periods (about 10 years of spore production), then ...

'' 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 suban ...

'' (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 beiteliana ''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 ...

'' Mickel *''

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'' (Alderw.) Holub *'' Huperzia catharinae'' (Christ) Holub *'' Huperzia ceylanica'' (Spring) Rothm. *'' Huperzia chinensis'' (Christ) Ching *'' Huperzia chishuiensis'' X.Y.Wang & P.S.Wang *'' Huperzia continentalis'' Testo, A.Haines & A.V.Gilman *'' Huperzia crispata'' (Ching) Ching *'' Huperzia delavayi'' (Christ & Herter) Ching *'' Huperzia emeiensis'' (Ching & H.S.Kung) Ching & H.S.Kung *'' Huperzia erosa'' Beitel & W.H.Wagner *'' Huperzia erubescens'' (Brack.) Holub *'' Huperzia europaea'' Björk, provisionally accepted *'' 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'' (also called 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 7–1 ...

'' (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 myriophyllifolia'' (Hayata) Holub *'' 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 porophila'' (F.E.Lloyd & Underw.) Holub *'' Huperzia quasipolytrichoides'' (Hayata) Ching *'' Huperzia rubicaulis'' S.K.Wu & X.Cheng *'' Huperzia saururoides'' (Bory & D'Urv.) Rothm. *'' Huperzia selago'' (L.) Bernh. (including ''H. selago'' subsp. ''appressa'' - Appalachian firmoss) *'' Huperzia serrata'' (Thunb.) Trevis. *'' Huperzia shresthae'' Fraser-Jenk. *'' Huperzia somae'' (Hayata) Ching *'' Huperzia sprengeri'' (Nessel) Holub *'' Huperzia suberecta'' (Lowe) Tardieu *'' Huperzia subintegra'' (Hillebr.) Beitel & W.H.Wagner *'' Huperzia sumatrana'' (Alderw.) Holub *'' Huperzia sutchueniana'' (Herter) Ching *'' Huperzia tibetica'' (Ching) Ching *'' Huperzia yakusimensis'' (Herter) Holub *'' Huperzia zollingeri'' (Herter) Holub The following hybrids have been described: * ''Huperzia'' × ''bartleyi'' (Cusick) Holub * ''Huperzia'' × ''buttersii'' (Abbe) Kartesz & Gandhi * ''Huperzia'' × ''carlquistii'' Beitel & W.H.Wagner * ''Huperzia'' × ''gillettii'' Beitel & W.H.Wagner * ''Huperzia'' × ''josephbeitelii'' A.Haines * ''Huperzia'' × ''medeirosii'' Beitel & W.H.Wagner * ''Huperzia'' × ''protoporophila'' A.Haines * ''Huperzia'' × ''sulcinervia'' (Spring) Trevis.

Distribution and habitat

circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...

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