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Spikemoss Scientific classification Kingdom: Plantae Division: Lycopodiophyta Class: Lycopodiopsida Order:
Spikemoss
Scientific classification
Kingdom:
Plantae
Division:
Lycopodiophyta
Class:
Lycopodiopsida
Order:
Selaginellales
Prantl
Family:
Selaginellaceae
Willk.
Genus:
Selaginella
P Beauv

.

.

Species

See text

. . Species See text Curled up Selaginella tamariscina Wallace's Selaginella (Selaginella wallacei)

Curled up Selaginella tamariscina

. . Species See text Curled up Selaginella tamariscina Wallace's Selaginella (Selaginella wallacei)

Wallace's Selaginella (Selaginella wallacei)

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". 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

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 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]

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 and exarch xylem in centre. 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. The stems contain no pith.

Unusually for the lycopods, which generally have microphylls with a single

unbranched vein, the microphylls of Selaginella species contain a branched vascular trace. In Selaginella, each microphyll and sporophyll has a small scale-like outgrowth called a ligule at the base of the upper surface. [3]:7 The plants are heterosporous with spores of two different size classes, known as megaspores and microspores.

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

Generic division

Some scientists still place the Selaginellales in the class Lycopodiopsida (often misconstructed as "Lycopsida"). Some modern authors recognize three generic divisions of Selaginella:

Selaginella, Bryodesma Sojak 1992, and Lycopodioides Boehm 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.

The first major attempt to define and subdivide the group was by Palisot de Beauvois [4] 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 [5][6] 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.

Walton & Aston Classification

In the Manual of Pteridology, [7] the

following classification was used by Walton & Alston:

genus: Selaginella

classification was used by Walton & Alston: genus: Selaginella subgenus: Euselaginella group: selaginoides

subgenus: Euselaginella group: selaginoides

classification was used by Walton & Alston: genus: Selaginella subgenus: Euselaginella group: selaginoides

group: pygmaea pygmaea

group: uliginosa ( Ericetorum ) uliginosa (Ericetorum)

group: rupestris ( Tetragonostachys or Bryodesma ) rupestris (Tetragonostachys or Bryodesma)

subgenus: Stachygynandrum Stachygynandrum

series: Decumbentes Decumbentes

series: Ascendentes Ascendentes

series: Sarmentosae Sarmentosae

series: Caulescentes Caulescentes

series: Circinatae Circinatae

series: Articulatae Articulatae

subgenus: Homostachys Homostachys

subgenus: Heterostachys Heterostachys

However, this is now known to be highly paraphyletic in most of its groupings. Two recent classifications, employing modern methods of phylogenetic analysis, are as follows:

Weststrand & Korall, 2016 Classification [8]

genus: Selaginella

subgenus: Selaginella Selaginella

clade: "Rhizophoric clade" clade A[ 8 ] genus: Selaginella subgenus: Selaginella subgenus Rupestrae (= Bryodesma Sojak or

Selaginella clade: "Rhizophoric clade" clade A subgenus Rupestrae (= Bryodesma Sojak or
Selaginella clade: "Rhizophoric clade" clade A subgenus Rupestrae (= Bryodesma Sojak or

subgenus Rupestrae (=Bryodesma Sojak or Tetragonostachys Jermy)

subgenus Lepidophyllae ( Selaginella lepidophylla ) Lepidophyllae (Selaginella lepidophylla)

subgenus Gymnogynum Gymnogynum

subgenus Exaltatae Exaltatae

subgenus Ericetorum (= Ericetorum Jermy) Ericetorum (=Ericetorum Jermy)

Exaltatae subgenus Ericetorum (= Ericetorum Jermy) clade B subgenus Stachygynandrum Zhang & Zhou, 2015

clade B

subgenus Stachygynandrumsubgenus Ericetorum (= Ericetorum Jermy) clade B Zhang & Zhou, 2015 Classification [ 9 ] genus:

Zhang & Zhou, 2015 Classification [9]

genus: Selaginella

Zhou, 2015 Classification [ 9 ] genus: Selaginella subgenus: Selaginella Type: Selaginella selaginoides

subgenus: Selaginella Type: Selaginella

selaginoides (L.) P.Beauv. ex Mart. & Schrank

subgenus: Boreoselaginella Type: Boreoselaginella Type:

Selaginella sanguinolenta (L.) Spring

subgenus: Ericetorum Type: Selaginella Ericetorum Type: Selaginella

uliginosa (Labill.) Spring

section: Lyallia Type: Selaginella Lyallia Type: Selaginella

uliginosa (Labill.) Spring

section: Myosurus Type: Selaginella myosurus Alston Myosurus Type: Selaginella myosurus Alston

section: Megalosporarum Type: Megalosporarum Type:

Selaginella exaltata (Kunze) Spring

section: Articulatae Type: Articulatae Type:

Selaginella kraussiana (Kunze)

A.Braun

section: Homoeophyllae Type: Homoeophyllae Type:

Selaginella rupestris (L.) Spring

(= Bryodesma Sojak or Tetragonostachys Jermy) section: Lepidophyllae Type: Selaginella lepidophylla (Hook. &

(=Bryodesma Sojak or Tetragonostachys Jermy)

section: Lepidophyllae Type:

Selaginella lepidophylla (Hook. &

Grev.) Spring

subgenus: Pulviniella Type: Selaginella Pulviniella Type: Selaginella

pulvinata (Hook. & Grev.) Maxim

subgenus: Heterostachys Type: Heterostachys Type:

Selaginella heterostachys Baker

section: Oligomacrosporangiatae Type: Selaginella uncinata (Desv. ex

section: Oligomacrosporangiatae Type: Selaginella uncinata (Desv. ex

Poir.) Spring section: Auriculatae Type:

Poir.) Spring section: Auriculatae Type:

Selaginella douglasii (Hook. &

Grev.) Spring

section: Homostachys Type: : Homostachys Type: :

Selaginella helvetica (L.) Link

section: Tetragonostachyae Type: Tetragonostachyae Type:

Selaginella proniflora (L.) Baker

section: Heterostachys Type: Heterostachys Type:

Selaginella brachystachya (Hook. &

Grev.) Spring

Type: Selaginella brachystachya (Hook. & Grev.) Spring subgenus: Stachygynandrum Type: Selaginella flabellata

subgenus: Stachygynandrum Type:

Selaginella flabellata (L.) Spring

section: Plagiophyllae Type: Plagiophyllae Type:

Selaginella biformis A.Braun ex

Kuhn

section: Circinatae Type: Selaginella Circinatae Type: Selaginella

involvens (Sw.) Spring

section: Heterophyllae Type: Heterophyllae Type:

Selaginella flexuosa Spring

section: Austroamericanae Type: Austroamericanae Type:

Selaginella hartwegiana Spring

section: Pallescentes Type: S. Pallescentes Type: S.

pallescens (C.Presl) Spring

section: Proceres Type: Selaginella oaxacana Spring Proceres Type: Selaginella oaxacana Spring

section: Ascendentes Type: Ascendentes Type:

Selaginella alopecuroides Baker

Species

Selaginella selaginoides

Selaginella willdenowii is known for its iridescent colours

There are about 750 known species of Selaginella, [10] showing a wide range of characters; the genus is overdue for a revision which might include subdivision into several genera. Better-known spikemosses include:

plant, dinosaur plant, and flower of stone (Chihuahuan Desert of North America)

spikemoss

Selaginella plana - Asian spikemoss Selaginella plana - Asian spikemoss

(Tropical Asia)

Selaginella rupestris - rock spikemoss, Selaginella rupestris - rock spikemoss,

festoon pine, and northern Selaginella (eastern North America)

Selaginella uncinata - peacock moss, Selaginella uncinata - peacock moss,

peacock spikemoss, blue spikemoss

Selaginella underwoodii Hieron. - (west Selaginella underwoodii Hieron. - (west

Texas to Wyoming and west into Arizona)

spikemoss, peacock fern (southeast Asia)

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:

S. kraussiana: golden clubmoss : golden clubmoss

S. moellendorffii: gemmiferous : gemmiferous

spikemoss

S. erythropus : red selaginella or ruby-red

S.

erythropus: red selaginella or ruby-red

spikemoss

S. uncinata : peacock moss

S.

uncinata: peacock moss

S. lepidophylla : resurrection plant

S.

lepidophylla: resurrection plant

S. braunii : arborvitae fern

S.

braunii: arborvitae fern

References

2009-04-08.

3. Stace, C. A. (2010). New Flora of the

British Isles (Third ed.). Cambridge, U.K.:

Cambridge University Press. ISBN 9780521707725.

4. Palisot de Beauvois (1805): Prodrome

des cinquième et sixième familles de l'Æthéogamie, les mousses, les lycopodes.

5. Korall, P. & Kenrick, P. (2002),

"Phylogenetic relationships in Selaginellaceae based on rbcL sequences", American Journal of Botany, 89 (3): 506–17, doi:10.3732/ajb.89.3.506 , PMID 21665649

6. Korall & Kenrick (2004): The

phylogenetic history of Selaginellaceae based on DNA sequences from the plastid and nucleus: extreme substitution rates

and rate heterogeneity. Molecular Phylogenetics and Evolution, Volume 31, Issue 3, June 2004, Pages 852-864

7. Verdoorn, F., ed. (1938): Manual of

Pteridology: J. Walton and A. H. G. Alston, Lycopodinae, pp. 500-506. Martinus Nijhoff, The Hague. 640pp, HB.

8. Weststrand, Stina; Korall, Petra (2016),

9. Zhou, Xin-Mao; Zhang, Li-Bing (2015), "A

classification of Selaginella

(Selaginellaceae) based on molecular (chloroplast and nuclear), macromorphological, and spore features", Taxon, 64 (6): 1117–1140,

External links

Media related to Selaginella at Wikimedia Commons Selaginella at Wikimedia Commons

Data related to Selaginella at Wikispecies Selaginella at Wikispecies