Selaginella

Last updated

Spikemoss
Temporal range: Late Carboniferous–Recent
Selaginella-sp.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Lycophytes
Class: Lycopodiopsida
Order: Selaginellales
Prantl
Family: Selaginellaceae
Willk.
Genus: Selaginella
P. Beauv.
Type species
Selaginella selaginoides
Species

See text.

Synonyms
  • BryodesmaSoják 1992
  • CarpolepidiumPalisot de Beauvois 1805
  • DidiclisPalisot de Beauvois 1803
  • DiplostachyumPalisot de Beauvois 1805
  • GymnogynumPalisot de Beauvois 1804
  • HeterophyllaeSpring 1840
  • HeterophylliumHieronymus ex Börner 1912
  • HomoeophyllaeSpring 1840
  • HomostachysWarburg 1900
  • HypopterygiopsisSakurai 1943
  • LycopodioidesBoehm. 1760 ex Kuntze 1891
  • MirmauAdanson 1763
  • PolycoccaHill 1773 nom. superfl.
  • SelaginoidesSéguier 1754 nom. rej.
  • SelagoBrowne 1756 nom. ill.
  • StachygynandrumPalisot de Beauvois 1804 nom. rej.
  • TrispermiumHill 1773
Curled up Selaginella tamariscina Spikemoss Selaginella Tamariscina curled up.jpg
Curled up Selaginella tamariscina
Wallace's Selaginella (Selaginella wallacei) Selaginella-wallacei.JPG
Wallace's Selaginella (Selaginella wallacei)

Selaginella, also known as spikemosses or lesser clubmosses is a genus of lycophyte. It is usually treated as the only genus in the family Selaginellaceae, with over 750 known species.

Contents

This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having spores of two types. They are sometimes included in an informal paraphyletic group called the "fern allies". The species S. moellendorffii is an important model organism. Its genome has been sequenced by the United States Department of Energy's Joint Genome Institute. [1] The name Selaginella was erected by Palisot de Beauvois solely for the species Selaginella selaginoides , which turns out (with the closely related Selaginella deflexa ) to be a clade that is sister to all other Selaginellas, so any definitive subdivision of the species into separate genera leaves two taxa in Selaginella, with the hundreds of other species in new or resurrected genera.

Selaginella occurs mostly in the tropical regions of the world, with a handful of species to be found in the arctic-alpine zones of both hemispheres. [2] Fossils assignable to the modern genus are known spanning over 300 million years from the Late Carboniferous to the present. [3]

Description

Selaginella species are creeping or ascendant plants with simple, scale-like leaves (microphylls) on branching stems from which roots also arise. The stems are aerial, horizontally creeping on the substratum (as in Selaginella kraussiana ), sub-erect ( Selaginella trachyphylla ) or erect (as in Selaginella erythropus ). The vascular steles are polystelic protosteles. Stem section shows the presence of more than two protosteles. Each stele is made up of diarch (having two strands of xylem [4] ) and exarch (growing outward in) xylems.[ citation needed ] The steles are connected with the cortex by means of many tube-like structures called trabeculae, which are modified endodermal cells with casparian strips on their lateral walls.[ citation needed ] The stems contain no pith.[ citation needed ]

In Selaginella, each microphyll and sporophyll has a small scale-like outgrowth called a ligule at the base of the upper surface. [5] :7 The plants are heterosporous with spores of two different size classes, known as megaspores and microspores. [6]

Unusual for the lycopods, which nearly always have microphylls with a single unbranched vein, the microphylls of a few Selaginella species contain a branched vascular trace. [7]

Under dry conditions, some species of Selaginella can survive dehydration. In this state, they may roll up into brown balls and be uprooted, but can rehydrate under moist conditions, become green again and resume growth. This phenomenon is known as poikilohydry, and poikilohydric plants such as Selaginella bryopteris are sometimes referred to as resurrection plants. [8]

There is no evidence of whole genome duplication in Selaginella's evolutionary history. Instead they have gone through tandem gene duplications, which is particularly noticeable in genes relevant for desiccation tolerance. [9] [ jargon ]

Their chloroplasts are missing about two-thirds of their plastidial tRNA genes, which are instead found in the genome of the nucleus. [10] The genus is unique among vascular plants in having species with monoplastidic cells, single giant chloroplasts, located mostly in their dorsal epidermal cells, but also in the upper mesophyll of some species. This appears to be a derived traits and an adaptation to low-light conditions, having originated at least twice. Cells with multiplastidic chloroplasts, more than ten chloroplasts per cell, are considered most basal, and are found in species exposed to more light. Oligoplastidic cells, cells with 3 to ten chloroplats, are more adated to weaker light, and the monoplastidic species being the most shade-loving forms. It is estimated that 70% of Selaginella species are monoplastidic. These receive just 0.4~2.1% of full sunlight, while species with multiple chloroplasts live in open places where they on average receive more than 40.5% of full sunlight. [11]

Taxonomy

Some scientists still place the Selaginellales in the class Lycopodiopsida (often misconstructed as "Lycopsida").[ citation needed ] [12] [13] Some modern authors recognize three generic divisions of Selaginella: Selaginella, BryodesmaSojak 1992, and LycopodioidesBoehm 1760. Lycopodioides would include the North American species S. apoda and S. eclipes, while Bryodesma would include S. rupestris (as Bryodesma rupestre). Stachygynandrum is also sometimes used to include the bulk of species.[ citation needed ]

The first major attempt to define and subdivide the group was by Palisot de Beauvois [14] in 1803–1805. He established the genus Selaginella as a monotypic genus, and placed the bulk of species in Stachygynandrum. Gymnogynum was another monotypic genus, but that name is superseded by his own earlier name of Didiclis. This turns out, today, to be a group of around 45–50 species also known as the Articulatae, since his genus Didiclis/Gymnogynum was based on Selaginella plumosa . He also described the genus Diplostachyum to include a group of species similar to Selaginella apoda. Spring inflated the genus Selaginella to hold all selaginelloid species four decades later.

Phylogenetic studies by Korall & Kenrick [15] [16] determined that the Euselaginella group, comprising solely the type species, Selaginella selaginoides and a closely related Hawaiian species, Selaginella deflexa , is a basal and anciently diverging sister to all other Selaginella species. Beyond this, their study split the remainder of species into two broad groups, one including the Bryodesma species, the Articulatae , section Ericetorum Jermy and others, and the other centered on the broad Stachygynandrum group.

In 2023, Zhou & Zhang suggested that the genus should be broken up into 19 different genera. [17]

Walton & Aston 1938 [18] Weststrand & Korall 2016 [19]
  • subgenus: Euselaginella
    • group: selaginoides
    • group: pygmaea
    • group: uliginosa (Ericetorum)
    • group: rupestris (Tetragonostachys or Bryodesma)
  • subgenus: Stachygynandrum
    • series: Decumbentes
    • series: Ascendentes
    • series: Sarmentosae
    • series: Caulescentes
    • series: Circinatae
    • series: Articulatae
  • subgenus: Homostachys
  • subgenus: Heterostachys
  • subgenus: Selaginella
  • clade: "Rhizophoric clade"
    • clade A
      • subgenus Rupestrae [Bryodesma Sojak or Tetragonostachys Jermy, S. section Homeophyllae]
      • subgenus Lepidophyllae [S. section Lepidophyllae]
      • subgenus Gymnogynum [S. section Articulatae]
      • subgenus Exaltatae [incl. S. section Megalosporum, S. section Myosurus]
      • subgenus Ericetorum [S. section Lyallia]
    • clade B
      • subgenus Stachygynandrum [incl. S. (Boreoselaginella), S. (Pulviniella), S. (Heterostachys)]

Zhang & Zhou, 2015 classification

[20]

Zhang & Zhou 2015 [20] Zhou & Zhang 2023 [17]
Selaginella

subgenus Selaginella

subgenus Boreoselaginella

subgenus

section Megalosporum

section Myosurus

section Lyallia

section Articulatae

section Lepidophyllae

section Homeophyllae

Gymnogynum

subgenus Pulviniella

subgenus

section Tetragonostachyae

section Heterostachys

section Auriculatae

section Homostachys

section

S. braunii subclade

S. willdenowii subclade

S. pennata subclade

S. pervillei subclade

S. siamensis subclade

S. delicatula subclade

Oligomacrosporangiatae
Heterostachys
subgenus

section Plagiophyllae

section Circinatae

section Ascendentes

section Proceres

section Pallescentes

section Austroamericanae

section Heterophyllae

Stachygynandrum
Selaginellaceae
Selaginoidoideae

SelaginoidesSéguier 1754

Boreoselaginelloideae

Boreoselaginella(Warburg 1900) Zhang & Zhou 2023

Gymnogynoideae

AfroselaginellaZhang & Zhou 2023

MegaloselaginellaZhang & Zhou 2023

Ericetorum(Jermy 1986) Zhang & Zhou 2023

GymnogynumPalisot de Beauvois 1804

LepidoselaginellaZhang & Zhou 2023

BryodesmaSoják 1992

Pulvinielloideae

Pulviniella(Zhang & Zhou 2015) Zhang & Zhou 2023

Sinoselaginelloideae

SinoselaginellaZhang & Zhou 2023

AustroselaginellaZhang & Zhou 2023

KoralliaZhang & Zhou 2023

Lycopodioidoideae

ValdespinoaZhang & Zhou 2023

LycopodioidesBoehm. 1760 ex Kuntze 1891

DidiclisPalisot de Beauvois ex Mirb. 1802

HypopterygiopsisSakurai 1943

Selaginelloideae

ChuselaginellaZhang & Zhou 2023

KungiselaginellaZhang & Zhou 2023

Selaginella

section ProceresSpring 1850

section PallescentesZhang & Zhou 2015

section AustroamericanaeZhang & Zhou 2015

section HeterophyllaeSpring 1840

Species

Selaginella canaliculata Selaginella canaliculata.jpeg
Selaginella canaliculata
Selaginella selaginoides Selaginella selaginoides - Niitvalja bog.jpg
Selaginella selaginoides
Selaginella willdenowii is known for its iridescent colours Iridescent fern.jpg
Selaginella willdenowii is known for its iridescent colours

There are about 750 known species of Selaginella. [21] They show a wide range of characters; the genus is overdue for a revision which might include subdivision into several genera.[ citation needed ] Species of spikemoss include:

A few species of Selaginella are desert plants known as "resurrection plants", because they curl up in a tight, brown or reddish ball during dry times, and uncurl and turn green in the presence of moisture. Other species are tropical forest plants that appear at first glance to be ferns.

Cultivation

A number of Selaginella species are popular plants for cultivation, mostly tropical species. Some of the species popularly cultivated and actively available commercially include:

Related Research Articles

<span class="mw-page-title-main">Lycopodiopsida</span> Class of vascular plants

Lycopodiopsida is a class of vascular plants also known as lycopsids, lycopods, or lycophytes. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching stems bearing simple leaves called microphylls and reproduce by means of spores borne in sporangia on the sides of the stems at the bases of the leaves. Although living species are small, during the Carboniferous, extinct tree-like forms (Lepidodendrales) formed huge forests that dominated the landscape and contributed to coal deposits.

<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>Selaginella lepidophylla</i> Species of spore-bearing plant

Selaginella lepidophylla, also known as a resurrection plant, is a species of desert plant in the spikemoss family (Selaginellaceae). It is native to the Chihuahuan Desert of the United States and Mexico. S. lepidophylla is renowned for its ability to survive almost complete desiccation. Resurrection plants are vascular rooted plants capable of surviving extreme desiccation, then resuming normal metabolic activity upon rehydration. The plant's hydro-responsive movements are governed by stem moisture content, tissue properties and a graded distribution of lignified cells affecting concentric stem stiffness and spiraling. During dry weather in its native habitat, its stems curl into a tight ball, uncurling only when exposed to moisture.

<i>Selaginella moellendorffii</i> Species of spore-bearing plant

Selaginella moellendorffii is a lycophyte that is an important model organism, especially in comparative genomics. S. moellendorffii is a member of an ancient vascular plant lineage that first appeared in the fossil record some 400 million years ago. They would later form a dominant part of the world's flora during the Carboniferous period. They have a number of unusual and/or "primitive" features, such as rudimentary leaves (microphylls), ubiquitous dichotomous branching, heterospory, and the ligule. As the earliest diverging group of modern vascular plants, they are essential to understanding the evolution of plants as a whole.

<i>Phlegmariurus phlegmaria</i> Species of fern

Phlegmariurus phlegmaria, synonym Huperzia phlegmaria, commonly known as either coarse tassel fern or common tassel fern, is an epiphytic species native to rainforests in Madagascar, some islands in the Indian Ocean, Asia, Australasia and many Pacific Islands. Phlegmariurus phlegmaria is commonly found in moist forests and rainforests at high altitudes, in and amongst mosses and other epiphytes. Members of the order Lycopodiales are commonly referred to as clubmosses.

<i>Selaginella apoda</i> Species of spore-bearing plant

Selaginella apoda, commonly known as meadow spikemoss, is a perennial lycophyte native to much of the eastern United States and parts of northeastern Mexico. The life cycle is the shortest of the genus Selaginella, as well as one of the shortest among the lycophytes. Selaginella apoda is found primarily in damp soils in habitats such as swamps, wet fields, open woods and along stream banks. Selaginella apoda presents the potential for case studies involving the plant's adaptability to environmental toxins. It is closely related to Selaginella eclipes and S. ludoviciana, with both of which it has been reported to form hybrids. This group is characterized by relatively flat strobili and large megasporophylls which occur in the same plane as the lateral leaves.

<i>Selaginella selaginoides</i> Species of spore-bearing plant

Selaginella selaginoides is a non-flowering plant of the spikemoss genus Selaginella with a wide distribution around the Northern Hemisphere. It resembles a moss in appearance but is a vascular plant belonging to the division Lycopodiophyta. It has a number of common names including lesser clubmoss, club spikemoss, northern spikemoss, low spikemoss and prickly mountain-moss. This plant has one close relative, Selaginella deflexa, native to Hawaii. These two plants form a small clade that is sister to all other Selaginella species.

Rose of Jericho or Jericho rose may refer to:

<i>Selaginella kraussiana</i> Species of clubmoss in the family Selaginellaceae

Selaginella kraussiana is a species of vascular plant in the family Selaginellaceae. It is referred to by the common names Krauss' spikemoss, Krauss's clubmoss, or African clubmoss, and is found naturally in parts of Sub-Saharan Africa and in Macaronesia. It is sometimes given the misnomer of “peacock fern”, due to its lacy leaf structure, despite having no relation to actual ferns; rather, it belongs to the very ancient lineage of plants known as the clubmosses.

<i>Selaginella uliginosa</i> Species of spikemoss

Selaginella uliginosa is a small plant in the spikemoss family Selaginellaceae which is endemic to Australia. An ancient and primitive plant, usually under 10 centimetres tall, it is often seen in sunny moist areas. The specific epithet uliginosa is from Latin, referring to the plant's preference for growing in swampy locations.

<i>Selaginella stellata</i> Species of spore-bearing plant

Selaginella stellata, also recognized by its common name, starry spikemoss or starry spike-moss, is a species of spikemoss of the family Selaginellaceae. It is a type of lycopod that grows naturally in Mexico and Central American countries like Guatemala and Belize and can also be found in the state of Hawaii.

<i>Selaginella willdenowii</i> Species of plant

Selaginella willdenowii is a species of vascular plant in the Selaginellaceae family. It is a spikemoss known by the common names Willdenow's spikemoss and peacock fern due to its iridescent blue leaves. Like other Selaginallales, it is fern ally and not a true fern.

The Melaniellaceae are a family of fungi in the division Basidiomycota and order of Doassansiales. The family contains 1 genera and 2 species. They have a distribution in south and south-east Asia.

<i>Selaginella australiensis</i> Species of spore-bearing plant

Selaginella australiensis is a plant in the spikemoss family Selaginellaceae endemic to northeastern Queensland. It grows in rainforest and closed forest from Cooktown to near Mission Beach, including the Atherton Tablelands. It is a low growing and much branched terrestrial plant inhabiting damp shady locations, typically along stream banks.

<i>Selaginella brisbanensis</i> Species of plant

Selaginella brisbanensis is a plant in the spikemoss family Selaginellaceae endemic to northeastern and southeastern Queensland. It grows in rainforest and wet sclerophyll forest in two very disjunct populations, one centred around Cairns and the other around Brisbane, some 1,400 km (870 mi) south. It is a terrestrial plant growing up to 20 cm (7.9 in) high.

<i>Selaginella <span style="font-style:normal;">×</span> neomexicana</i> Species of plant

Selaginella × neomexicana, commonly known New Mexican spikemoss, is a hybrid species of desert vascular plant in the spikemoss family Selaginellaceae. First described by William Ralph Maxon, it is found in the south-western United States.

<i>Selaginella deflexa</i> Species of spikemoss

Selaginella deflexa, commonly known as deflexed spikemoss, is a non-flowering species of plant in the spikemoss genus Selaginella and is in the subgenus of the same name. It is closely related to Selaginella selaginoides which is the only other member in its subgenus. It is endemic to Hawaii and grows primarily in the wet tropical biome from 1,050–1,500 m (3,440–4,920 ft). can be found on all major islands except for Lana'i. It is usually found growing in wet moss. Unlike most members of Selaginella,S. deflexa does not have rhizophores along its stem. It gets its name from its reflexed leaves which point outwards.

References

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