Huperzia

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Huperzia
Huperzia selago.jpg
Huperzia selago in Austria
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Clade: Lycophytes
Class: Lycopodiopsida
Order: Lycopodiales
Family: Lycopodiaceae
Subfamily: Huperzioideae
Genus: Huperzia
Bernh. [1]
Species

See text.

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. [2] 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 (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, [3] although not all have been formally transferred to other genera. [4] Other sources recognize only Huperzia, which then has about 340 species. [5]

Contents

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 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 gametophytes are not photosynthetic, but rather subterranean and mycorrhizal. [2]

The Flora of North America distinguishes Huperzia from the 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. [2]

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 . [1]

In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), Huperzia is placed in the subfamily Huperzioideae of the family Lycopodiaceae. [3] A 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 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: [6]

Huperzioideae

Phylloglossum

Huperzia

Phlegmariurus

The majority of the species formerly placed in a broadly defined Huperzia belong in Phlegmariurus. [6] Earlier, the Flora of North America had also separated Huperzia from Phegmariurus. [2] However, Phlegmariurus is difficult to separate morphologically, and others have preferred the one-genus division of the subfamily. [7] [5]

Species

The PPG I classification stated there were 25 species in the genus Huperzia. [3] As of January 2023, World Ferns listed 61 species, noting that "many species still need transfer into other split genera". [4]

The following hybrids have been described: [4]

Distribution and habitat

As circumscribed in the PPG I classification, Huperzia is distributed in temperate, arctic and alpine habitats, including mountains in tropical Asia. [4] Its species are terrestrial or grow on rocks. [2] Phlegmariurus is epiphytic, [2] and has a worldwide tropical distribution, [7] 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. [5]

Related Research Articles

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

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 recognizes three extant genera:

<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.

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

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<i>Arachniodes</i> Genus of ferns

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

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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 that reflects knowledge about plant relationships discovered through phylogenetic studies. In 2016, the group published a classification for extant pteridophytes, termed "PPG I". The paper had 94 authors.

<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

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<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">Lycopodioideae</span> Subfamily of spore-bearing plants

Lycopodioideae 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 Lycopodium in other classifications. Like all lycophytes, members of the Lycopodioideae reproduce by spores. The oldest fossils of modern members of the family date to the Early Cretaceous.

<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>Phlegmariurus squarrosus</i> Species of spore-bearing plant

Phlegmariurus squarrosus is a species of lycophyte in the family Lycopodiaceae. The genus Phlegmariurus is accepted in the Pteridophyte Phylogeny Group classification of 2016, but not in other classifications, which submerge the genus in Huperzia, with this species as Huperzia squarrosa. The species has a wide distribution from the west Indian Ocean, through tropical and subtropical Asia to eastern Australia and the Pacific.

<i>Phlegmariurus dentatus</i> Species of flowering plant

Phlegmariurus dentatus is a species of plant in the family Lycopodiaceae, endemic to the Portuguese archipelagos of the Azores and Madeira. In the Azores, it appears in sloping areas, woods, forests and sometimes on road sides, being less demanding in humidity than Huperzia suberecta. It is native to seven of the nine islands. In the archipelago of Madeira it only occurs on the island of Madeira.

References

  1. 1 2 "Huperzia Bernh.", The International Plant Names Index , retrieved 2020-09-16
  2. 1 2 3 4 5 6 Wagner Jr., Warren H. & Beitel, Joseph M., "Huperzia", in Flora of North America Editorial Committee (ed.), Flora of North America (online), eFloras.org, retrieved 2020-09-16
  3. 1 2 3 PPG I (2016), "A community-derived classification for extant lycophytes and ferns", Journal of Systematics and Evolution, 54 (6): 563–603, doi:10.1111/jse.12229, S2CID   39980610
  4. 1 2 3 4 Hassler, Michael (19 January 2023), "Huperzia", World Ferns. Synonymic Checklist and Distribution of Ferns and Lycophytes of the World, 14.7, retrieved 2023-01-22
  5. 1 2 3 "Huperzia Bernh.", Plants of the World Online, Royal Botanic Gardens, Kew, retrieved 2020-09-15
  6. 1 2 Field, Ashley R.; Testo, Weston; Bostock, Peter D.; Holtum, Joseph A. M. & Waycott, Michelle (2016), "Molecular phylogenetics and the morphology of the Lycopodiaceae subfamily Huperzioideae supports three genera: Huperzia, Phlegmariurus and Phylloglossum", Molecular Phylogenetics and Evolution, 94 (Pt B): 635–657, doi:10.1016/j.ympev.2015.09.024, PMID   26493224
  7. 1 2 Hassler, Michael (19 January 2023), "Phlegmariurus", World Ferns. Synonymic Checklist and Distribution of Ferns and Lycophytes of the World, 14.7, retrieved 2023-01-22
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