Editing Huperzia

{{Short description|Genus of vascular plants}}
{{Automatic taxobox
| image = Huperzia selago.jpg
| image_caption = ''Huperzia selago'' in Austria
| taxon = Huperzia
| authority = [[Johann Jacob Bernhardi|Bernh.]]<ref name=IPNI_30000333-2/>
| subdivision_ranks = Species
| subdivision = [[#Species|See text]].
}}

'''''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'''.<ref name=FNA/> 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 [[Circumscription (taxonomy)|circumscriptions]] of the genus were in use. In the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I), ''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,<ref name=PPGI/> although not all have been formally transferred to other genera.<ref name=CFLW_Hup/> Other sources recognize only ''Huperzia'', which then has about 340 species.<ref name=POWO_30000333-2/>

==Morphology==
The [[sporophyte]]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 [[Sporangium|sporangia]] 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]]s are not photosynthetic, but rather subterranean and [[mycorrhiza]]l.<ref name=FNA/>

The ''Flora of North America'' distinguishes ''Huperzia'' from the [[Epiphyte|epiphytic]] 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.<ref name=FNA/>

==Taxonomy==
The genus ''Huperzia'' was created by [[Johann Jakob Bernhardi]] in 1801. Bernhardi separated ''Huperzia'' from ''Lycopodium''. The [[type species]] is ''Lycopodium selago'' which became ''[[Huperzia selago]]''.<ref name=IPNI_30000333-2/>

In the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I), ''Huperzia'' is placed in the subfamily [[Huperzioideae]] of the family [[Lycopodiaceae]].<ref name=PPGI/> A [[Phylogenetics|phylogenetic]] study in 2016, employing both molecular and morphological data, concluded that either a one-genus or a three-genus division of the subfamily produced [[Monophyly|monophyletic]] taxa. The authors preferred the three-genus division, recognizing ''Huperzia'', ''[[Phlegmariurus]]'' and ''[[Phylloglossum]]''. Their preferred hypothesis for the relationships of the three genera was:<ref name=FielTestBostHolt16/>
{{clade
|label1=[[Huperzioideae]]
|1={{clade
   |1={{clade
      |1=''[[Phylloglossum]]''
      |2=''Huperzia''
      }}
   |2=''[[Phlegmariurus]]''
   }}
}}
The majority of the species formerly placed in a broadly defined ''Huperzia'' belong in ''Phlegmariurus''.<ref name=FielTestBostHolt16/> Earlier, the ''Flora of North America'' had also separated ''Huperzia'' from ''Phegmariurus''.<ref name=FNA/> However, ''Phlegmariurus'' is difficult to separate morphologically, and others have preferred the one-genus division of the subfamily.<ref name=CFLW_Phl/><ref name=POWO_30000333-2/>

===Species===
The PPG I classification stated there were 25 species in the genus ''Huperzia''.<ref name=PPGI/> {{As of|2024|June}}, ''World Ferns'' listed the following species, noting that "many species still need transfer into other split genera".<ref name=CFLW_Hup/>
{{Div col|colwidth=25em}}
*''[[Huperzia acicularis]]'' <small>Björk</small>, provisionally accepted
*''[[Huperzia appalachiana]]'' <small>Beitel & Mickel</small>
*''[[Huperzia archboldiana]]'' <small>(Nessel) Holub</small>
*''[[Huperzia arunachalensis]]'' <small>(D.D.Pant & P.S.Pandey) Fraser-Jenk.</small>
*''[[Huperzia asiatica]]'' <small>(Ching) N.Shrestha & X.C.Zhang</small>
*''[[Huperzia australiana]]'' <small>(Herter) Holub</small>
*''[[Huperzia beccarii]]'' <small>(Alderw.) Holub</small>
*''[[Huperzia beiteliana]]'' <small>Mickel</small>
*''[[Huperzia bucahwangensis]]'' <small>Ching</small>
*''[[Huperzia campestris]]'' <small>(Alderw.) Holub</small>
*''[[Huperzia catharinae]]'' <small>(Christ) Holub</small>
*''[[Huperzia ceylanica]]'' <small>(Spring) Rothm.</small>
*''[[Huperzia chinensis]]'' <small>(Christ) Ching</small>
*''[[Huperzia chishuiensis]]'' <small>X.Y.Wang & P.S.Wang</small>
*''[[Huperzia continentalis]]'' <small>Testo, A.Haines & A.V.Gilman</small>
*''[[Huperzia crispata]]'' <small>(Ching) Ching</small>
*''[[Huperzia delavayi]]'' <small>(Christ & Herter) Ching</small>
*''[[Huperzia emeiensis]]'' <small>(Ching & H.S.Kung) Ching & H.S.Kung</small>
*''[[Huperzia erosa]]'' <small>Beitel & W.H.Wagner</small>
*''[[Huperzia erubescens]]'' <small>(Brack.) Holub</small>
*''[[Huperzia europaea]]'' <small>Björk</small>, provisionally accepted
*''[[Huperzia everettii]]'' <small>(Herter) Holub</small>
*''[[Huperzia fuegiana]]'' <small>(Roiv.) Holub</small>
*''[[Huperzia gedeana]]'' <small>(Alderw.) Holub</small>
*''[[Huperzia goliathensis]]'' <small>(Alderw.) Holub</small>
*''[[Huperzia haleakalae]]'' <small>(Brack.) Holub</small>
*''[[Huperzia herteriana]]'' <small>(Kümmerle) T.Sen & U.Sen</small>
*''[[Huperzia javanica]]'' <small>(Sw.) Fraser-Jenk.</small>
*''[[Huperzia jejuensis]]'' <small>B.Y.Sun & J.Lim</small>
*''[[Huperzia kangdingensis]]'' <small>(Ching) Ching</small>
*''[[Huperzia kunmingensis]]'' <small>Ching</small>
*''[[Huperzia laipoensis]]'' <small>Ching</small>
*''[[Huperzia lajouensis]]'' <small>Ching</small>
*''[[Huperzia leishanensis]]'' <small>X.Y.Wang</small>
*''[[Huperzia liangshanica]]'' <small>(H.S.Kung) Ching & H.S.Kung</small>
*''[[Huperzia lucidula]]'' <small>(Michx.) Trevis.</small>
*''[[Huperzia medogensis]]'' <small>Ching & Y.X.Lin</small>
*''[[Huperzia meghalaica]]'' <small>Fraser-Jenk.</small>
*''[[Huperzia miniata]]'' <small>(Spring) Trevis.</small>
*''[[Huperzia minima]]'' <small>(Herter) Holub</small>
*''[[Huperzia miyoshiana]]'' <small>(Makino) Ching</small>
*''[[Huperzia muscicola]]'' <small>W.M.Chu</small>
*''[[Huperzia myriophyllifolia]]'' <small>(Hayata) Holub</small>
*''[[Huperzia nanchuanensis]]'' <small>(Ching & H.S.Kung) Ching & H.S.Kung</small>
*''[[Huperzia nanlingensis]]'' <small>Shrestha, F.W.Xing, X.P.Qi, Y.H.Yan & X.C.Zhang</small>
*''[[Huperzia occidentalis]]'' <small>(Clute) Kartesz & Gandhi</small>
*''[[Huperzia porophila]]'' <small>(F.E.Lloyd & Underw.) Holub</small>
*''[[Huperzia quasipolytrichoides]]'' <small>(Hayata) Ching</small>
*''[[Huperzia rubicaulis]]'' <small>S.K.Wu & X.Cheng</small>
*''[[Huperzia saururoides]]'' <small>(Bory & D'Urv.) Rothm.</small>
*''[[Huperzia selago]]'' <small>(L.) Bernh.</small> (including [[Huperzia appressa|''H.&nbsp;selago'' subsp. ''appressa'']] - Appalachian firmoss)
*''[[Huperzia serrata]]'' <small>(Thunb.) Trevis.</small>
*''[[Huperzia shresthae]]'' <small>Fraser-Jenk.</small>
*''[[Huperzia somae]]'' <small>(Hayata) Ching</small>
*''[[Huperzia sprengeri]]'' <small>(Nessel) Holub</small>
*''[[Huperzia suberecta]]'' <small>(Lowe) Tardieu</small>
*''[[Huperzia subintegra]]'' <small>(Hillebr.) Beitel & W.H.Wagner</small>
*''[[Huperzia sumatrana]]'' <small>(Alderw.) Holub</small>
*''[[Huperzia sutchueniana]]'' <small>(Herter) Ching</small>
*''[[Huperzia tibetica]]'' <small>(Ching) Ching</small>
*''[[Huperzia yakusimensis]]'' <small>(Herter) Holub</small>
*''[[Huperzia zollingeri]]'' <small>(Herter) Holub</small>
{{Div col end}}

The following hybrids have been described:<ref name=CFLW_Hup/>
*[[Huperzia × bartleyi|''Huperzia'' × ''bartleyi'']] <small>(Cusick) Holub</small>
*[[Huperzia × buttersii|''Huperzia'' × ''buttersii'']] <small>(Abbe) Kartesz & Gandhi</small>
*[[Huperzia × carlquistii|''Huperzia'' × ''carlquistii'']] <small>Beitel & W.H.Wagner</small>
*[[Huperzia × gillettii|''Huperzia'' × ''gillettii'']] <small>Beitel & W.H.Wagner</small>
*[[Huperzia × josephbeitelii|''Huperzia'' × ''josephbeitelii'']] <small>A.Haines</small>
*[[Huperzia × medeirosii|''Huperzia'' × ''medeirosii'']] <small>Beitel & W.H.Wagner</small>
*[[Huperzia × protoporophila|''Huperzia'' × ''protoporophila'']] <small>A.Haines</small>
*[[Huperzia × sulcinervia|''Huperzia'' × ''sulcinervia'']] <small>(Spring) Trevis.</small>

==Distribution and habitat==
As [[Circumscription (taxonomy)|circumscribed]] in the PPG I classification, ''Huperzia'' is distributed in temperate, arctic and alpine habitats, including mountains in tropical Asia.<ref name=CFLW_Hup/> Its species are terrestrial or grow on rocks.<ref name=FNA/> ''Phlegmariurus'' is [[Epiphyte|epiphytic]],<ref name=FNA/> and has a worldwide tropical distribution,<ref name=CFLW_Phl/> 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.<ref name=POWO_30000333-2/>

== References ==
{{Reflist|refs=
<ref name=CFLW_Hup>{{cite WF |taxon=Huperzia |access-date=2024-06-02}}</ref>

<ref name=CFLW_Phl>{{citation |last=Hassler |first=Michael |contribution=Phlegmariurus |title=World Ferns. Synonymic Checklist and Distribution of Ferns and Lycophytes of the World |url=https://www.worldplants.de/ferns/ |at=14.7 |date=19 January 2023 |accessdate=2023-01-22}}</ref>

<ref name=FielTestBostHolt16>{{Citation |last1=Field |first1=Ashley R. |last2=Testo |first2=Weston |last3=Bostock |first3=Peter D. |last4=Holtum |first4=Joseph A. M. |last5=Waycott |first5=Michelle |date=2016 |title=Molecular phylogenetics and the morphology of the Lycopodiaceae subfamily Huperzioideae supports three genera: Huperzia, Phlegmariurus and Phylloglossum |journal=Molecular Phylogenetics and Evolution |volume=94 |issue=Pt B |pages=635–657 |doi=10.1016/j.ympev.2015.09.024 |pmid=26493224 |bibcode=2016MolPE..94..635F |name-list-style=amp }}</ref>

<ref name=FNA>{{Citation |first1=Warren H. |last1=Wagner Jr. |first2=Joseph M. |last2=Beitel |contribution=Huperzia |contribution-url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=115890 |accessdate=2020-09-16 |name-list-style=amp |editor-last=Flora of North America Editorial Committee |title=Flora of North America ''(online)'' |publisher=eFloras.org }}</ref>

<ref name=IPNI_30000333-2>{{citation |title=''Huperzia'' Bernh. |work=[[International Plant Names Index|The International Plant Names Index]] |url=https://www.ipni.org/n/30000333-2 |accessdate=2020-09-16 }}</ref>

<ref name=POWO_30000333-2>{{citation |title=''Huperzia'' Bernh. |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30000333-2 |accessdate=2020-09-15 }}</ref>

<ref name=PPGI>{{citation |author=PPG I |year=2016 |title=A community-derived classification for extant lycophytes and ferns |journal=Journal of Systematics and Evolution |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229|s2cid=39980610 |doi-access=free }}</ref>
}}

== External links ==
*[https://web.archive.org/web/20110610111056/http://botit.botany.wisc.edu/courses/img/bot/401/Lycophyta_(Microphyllophyta)/Lycopodiaceae/Huperzia/ Picture gallery]

{{Taxonbar|from=Q2111150}}

[[Category:Huperzia| ]]
[[Category:Lycophyte genera]]