Marchantiophyta

From Wikipedia, the free encyclopedia

The Marchantiophyta /mrkntift/ are a division of non-

vascular land plants commonly referred to as hepatics or liverworts. Liverworts
Like mosses and hornworts, they have a gametophyte-dominant life Temporal range: 4720 Ma
cycle, in which cells of the plant carry only a single set of genetic
Pre OS D C P T J K Pg N
information.
Mid-Ordovician[1] to present
It is estimated that there are about 9000 species of liverworts. Some of
the more familiar species grow as a flattened leafless thallus, but most
species are leafy with a form very much like a flattened moss. Leafy
species can be distinguished from the apparently similar mosses on the
basis of a number of features, including their single-celled rhizoids.
Leafy liverworts also differ from most (but not all) mosses in that their
leaves never have a costa (present in many mosses) and may bear
marginal cilia (very rare in mosses). Other differences are not universal
for all mosses and liverworts, but the occurrence of leaves arranged in
three ranks, the presence of deep lobes or segmented leaves, or a lack
of clearly differentiated stem and leaves all point to the plant being a
liverwort.

Liverworts are typically small, usually from 220 mm wide with

individual plants less than 10 cm long, and are therefore often
overlooked. However, certain species may cover large patches of
ground, rocks, trees or any other reasonably firm substrate on which
they occur. They are distributed globally in almost every available "Hepaticae" from Ernst Haeckel's
habitat, most often in humid locations although there are desert and Kunstformen der Natur, 1904
Arctic species as well. Some species can be a nuisance in shady
greenhouses or a weed in gardens.[4] Scientific classification
Kingdom: Plantae
Division: Marchantiophyta
Contents Stotler & Stotl.-Crand.,
1977 [2] emend. 2000 [3]
1 Physical characteristics
1.1 Description Classes and Orders
1.2 Life cycle
1.2.1 Asexual reproduction
2 Ecology Haplomitriopsida
3 Classification Haplomitriales
3.1 Relationship to other plants Treubiales
3.2 Internal classification
4 Economic importance Marchantiopsida
5 Gallery Blasiales
6 See also Neohodgsoniales
7 References
8 External links Sphaerocarpales
Lunulariales
Marchantiales
Physical characteristics Jungermanniopsida
Pelliales
Description Pallaviciniales
Fossombroniales
Most liverworts are small, measuring from 220 millimetres (0.08 Pleuroziales
0.8 in) wide with individual plants less than 10 centimetres (4 in) Metzgeriales
long,[5] so they are often overlooked. The most familiar liverworts
consist of a prostrate, flattened, ribbon-like or branching structure Porellales
called a thallus (plant body); these liverworts are termed thallose Ptilidiales
liverworts. However, most liverworts produce flattened stems with Jungermanniales
overlapping scales or leaves in two or more ranks, the middle rank is
often conspicuously different from the outer ranks; these are called
leafy liverworts or scale liverworts.[6][7] (See the gallery below for examples.)

Liverworts can most reliably be distinguished from the apparently

similar mosses by their single-celled rhizoids.[8] Other differences are
not universal for all mosses and all liverworts;[7] but the lack of clearly
differentiated stem and leaves in thallose species, or in leafy species the
presence of deeply lobed or segmented leaves and the presence of
leaves arranged in three ranks, all point to the plant being a
liverwort.[9][10] Unlike any other embryophytes, most liverworts
contain unique membrane-bound oil bodies containing isoprenoids in at
least some of their cells, lipid droplets in the cytoplasm of all other
A thallose liverwort, Lunularia cruciata plants being unenclosed.[11] The overall physical similarity of some
mosses and leafy liverworts means that confirmation of the
identification of some groups can be performed with certainty only with
the aid of microscopy or an experienced bryologist.

Liverworts have a gametophyte-dominant life cycle, with the sporophyte dependent on the gametophyte.[11]
Cells in a typical liverwort plant each contain only a single set of genetic information, so the plant's cells are
haploid for the majority of its life cycle. This contrasts sharply with the pattern exhibited by nearly all animals
and by most other plants. In the more familiar seed plants, the haploid generation is represented only by the tiny
pollen and the ovule, while the diploid generation is the familiar tree or other plant.[12] Another unusual feature
of the liverwort life cycle is that sporophytes (i.e. the diploid body) are very short-lived, withering away not
long after releasing spores.[13] Even in other bryophytes, the sporophyte is persistent and disperses spores over
an extended period.

Life cycle

The life of a liverwort starts from the germination of a

haploid spore to produce a protonema, which is either a
mass of thread-like filaments or else a flattened
thallus.[14][15] The protonema is a transitory stage in the
life of a liverwort, from which will grow the mature
gametophore ("gamete-bearer") plant that produces the sex
organs. The male organs are known as antheridia (singular:
antheridium) and produce the sperm cells. Clusters of
antheridia are enclosed by a protective layer of cells called
the perigonium (plural: perigonia). As in other land
plants, the female organs are known as archegonia
(singular: archegonium) and are protected by the thin
surrounding perichaetum (plural: perichaeta).[7] Each
archegonium has a slender hollow tube, the "neck", down
which the sperm swim to reach the egg cell.

Liverwort species may be either dioicous or monoicous. In

dioicous liverworts, female and male sex organs are borne Life cycle of a Marchantia-like liverwort
on different and separate gametophyte plants. In
monoicous liverworts, the two kinds of reproductive structures are borne on different branches of the same
plant.[16] In either case, the sperm must move from the antheridia where they are produced to the archegonium
where the eggs are held. The sperm of liverworts is biflagellate, i.e. they have two tail-like flagellae that enable
them to swim short distances,[17] provided that at least a thin film of water is present. Their journey may be
assisted by the splashing of raindrops. In 2008, Japanese researchers discovered that some liverworts are able to
fire sperm-containing water up to 15 cm in the air, enabling them to fertilize female plants growing more than a
metre from the nearest male.[18]

When sperm reach the archegonia, fertilisation occurs, leading to the production of a diploid sporophyte. After
fertilisation, the immature sporophyte within the archegonium develops three distinct regions: (1) a foot, which
both anchors the sporophyte in place and receives nutrients from its "mother" plant, (2) a spherical or
ellipsoidal capsule, inside which the spores will be produced for dispersing to new locations, and (3) a seta
(stalk) which lies between the other two regions and connects them.[17] When the sporophyte has developed all
three regions, the seta elongates, pushing its way out of the archegonium and rupturing it. While the foot
remains anchored within the parent plant, the capsule is forced out by the seta and is extended away from the
plant and into the air. Within the capsule, cells divide to produce both elater cells and spore-producing cells.
The elaters are spring-like, and will push open the wall of the capsule to scatter themselves when the capsule
bursts. The spore-producing cells will undergo meiosis to form haploid spores to disperse, upon which point the
life cycle can start again.

Asexual reproduction

Some liverworts are capable of asexual reproduction; in bryophytes in general "it would almost be true to say
that vegetative reproduction is the rule and not the exception."[19] For example, in Riccia, when the older parts
of the forked thalli die, the younger tips become separate individuals.[19]

Some thallose liverworts such as Marchantia polymorpha and Lunularia cruciata produce small disc-shaped
gemmae in shallow cups.[20] Marchantia gemmae can be dispersed up to 120 cm by rain splashing into the
cups.[21] In Metzgeria, gemmae grow at thallus margins.[22] Marchantia polymorpha is a common weed in
greenhouses, often covering the entire surface of containers;[23]:230 gemma dispersal is the "primary
mechanism by which liverwort spreads throughout a nursery or greenhouse."[23]:231

Ecology
Today, liverworts can be found in many ecosystems across the planet except the sea and excessively dry
environments, or those exposed to high levels of direct solar radiation.[24] As with most groups of living plants,
they are most common (both in numbers and species) in moist tropical areas.[25] Liverworts are more
commonly found in moderate to deep shade, though desert species may tolerate direct sunlight and periods of
total desiccation.

Classification
Relationship to other plants

Traditionally, the liverworts were grouped together with other bryophytes (mosses and hornworts) in the
Division Bryophyta, within which the liverworts made up the class Hepaticae (also called
Marchantiopsida).[7][26] However, since this grouping makes the Bryophyta paraphyletic, the liverworts are
now usually given their own division.[27] The use of the division name Bryophyta sensu lato is still found in the
literature, but more frequently the Bryophyta now is used in a restricted sense to include only the mosses.
 Two hypotheses on thephylogeny of land plants (embryophyta).

Another reason that liverworts are now classified separately is that they appear to have diverged from all other
embryophyte plants near the beginning of their evolution. The strongest line of supporting evidence is that
liverworts are the only living group of land plants that do not have stomata on the sporophyte generation.[28]
Among the earliest fossils believed to be liverworts are compression fossils of Pallaviciniites from the Upper
Devonian of New York.[29] These fossils resemble modern species in the Metzgeriales.[30] Another Devonian
fossil called Protosalvinia also looks like a liverwort, but its relationship to other plants is still uncertain, so it
may not belong to the Marchantiophyta. In 2007, the oldest fossils assignable at that time to the liverworts were
announced, Metzgeriothallus sharonae from the Givetian (Middle Devonian) of New York, United States.[31]
However, in 2010, five different types of fossilized liverwort spores were found in Argentina, dating to the
much earlier Middle Ordovician, around 470 million years ago.[1][32]

Internal classification

Bryologists classify liverworts in the division Marchantiophyta. This divisional name is based on the name of
the most universally recognized liverwort genus Marchantia.[33] In addition to this taxon-based name, the
liverworts are often called Hepaticophyta. This name is derived from their common Latin name as Latin was
the language in which botanists published their descriptions of species. This name has led to some confusion,
partly because it appears to be a taxon-based name derived from the genus Hepatica which is actually a
flowering plant of the buttercup family Ranunculaceae. In addition, the name Hepaticophyta is frequently
misspelled in textbooks as Hepatophyta, which only adds to the confusion.

Although there is no consensus among bryologists as to the classification of liverworts above family rank,[34]
the Marchantiophyta may be subdivided into three classes:[35][36][37][38]

The Jungermanniopsida includes the two orders Metzgeriales (simple thalloids) and Jungermanniales
(leafy liverworts).
The Marchantiopsida includes the three orders Marchantiales (complex-thallus liverworts), and
Sphaerocarpales (bottle hepatics), as well as the Blasiales (previously placed among the
Metzgeriales).[35][39] It also includes the problematic genus Monoclea, which is sometimes placed in its
own order Monocleales.[40]
A third class, the Haplomitriopsida is newly recognized as a basal sister group to the other
liverworts;[38] it comprises the genera Haplomitrium, Treubia, and Apotreubia.

Marchantiophyta
Haplomitriales
The diagram at right summarizes a Haplomitriopsida
portion of a 2006 cladistic analysis of
Treubiales
liverworts based upon three
chloroplast genes, one nuclear gene,
Blasiales
and one mitochondrial gene.[35]
Marchantiopsida
Sphaerocarpales

Marchantiales
Jungermanniopsida
Metzgeriales (part)
 Jungermanniales

Metzgeriales (part)

An updated classification by Sderstrm et al. 2016[41]

Marchantiophyta Stotler & Crandall-Stotler 2000

Haplomitriopsida Stotler & Crandall-Stotler 1977
Haplomitriales Hamlin 1972
Treubiales Schljakov 1972
Marchantiopsida Cronquist, Takhtajan & Zimmermann 1966
Blasiidae He-Nygrn et al. 2006
Blasiales Stotler & Crandall-Stotler 2000
Marchantiidae Engler 1893 sensu He-Nygrn et al. 2006
Neohodgsoniales Long 2006
Sphaerocarpales Cavers 1910 (bottle liverworts)
Lunulariales Long 2006
Marchantiales Limpricht 1877 (complex thalloids)
Jungermanniopsida Stotler & Crandall-Stotler 1977
Pelliidae He-Nygrn et al. 2006
Pelliales He-Nygrn et al. 2006
Pallaviciniales Frey & Stech 2005
Fossombroniales Schljakov 1972
Metzgeriidae Bartholomew-Began 1990
Pleuroziales Schljakov 1972
Metzgeriales Chalaud 1930
Jungermanniidae Engler 1893 (leafy liverworts)
Porellales Schljakov 1972
Ptilidiales Schljakov 1972
Jungermanniales von Klinggrf f 1858

It is estimated that there are about 9000 species of liverworts, at least 85% of which belong to the leafy
group.[3][42] Despite that fact, no liverwort genomes have been sequenced to date and only few genes identified
and characterized.[43]

Economic importance
In ancient times, it was believed that liverworts cured diseases of the liver, hence the name.[44] In Old English,
the word liverwort literally means liver plant.[45] This probably stemmed from the superficial appearance of
some thalloid liverworts (which resemble a liver in outline), and led to the common name of the group as
hepatics, from the Latin word hpaticus for "belonging to the liver". An unrelated flowering plant, Hepatica, is
sometimes also referred to as liverwort because it was once also used in treating diseases of the liver. This
archaic relationship of plant form to function was based in the "Doctrine of Signatures".[46]

Liverworts have little direct economic importance today. Their greatest impact is indirect, through the reduction
of erosion along streambanks, their collection and retention of water in tropical forests, and the formation of
soil crusts in deserts and polar regions. However, a few species are used by humans directly. A few species,
such as Riccia fluitans, are aquatic thallose liverworts sold for use in aquariums. Their thin, slender branches
float on the water's surface and provide habitat for both small invertebrates and the fish that feed on them.

Gallery
A small collection of images showing liverwort structure and diversity:
 Marchantia The archegonium of A sporophyte emerging Porella platyphylla
polymorpha, with Porella. from its archegonium. clump growing on a
antheridial and tree.
archegonial stalks.

Pellia epiphylla, Plagiochila Riccia fluitans, an Conocephalum

growing on moist soil. asplenioides, a leafy aquatic thallose conicum, a large
liverwort. liverwort. thallose liverwort.

See also
Bryophyte
Embryophyte

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External links
Picture Gallery of Mosses & Liverworts
Liverwort structure in pictures
Liverwort classification scheme
LiToL: Assembling the Liverwort Tree of Life (note: for 500,000 million years ago read "480 million
years ago".)
Inter-relationships of Mosses, Liverworts, and Hornworts
Moss-Liverwort connection
Additional information on Liverworts
Liverworts

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