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Marchantiophyta
The Marchantiophyta /mɑːrˌkæntiˈɒfɪtə/ ( listen) are a division
of non-vascular land plants commonly referred to as hepatics or Liverworts
liverworts. Like mosses and hornworts, they have a Temporal range: 472–0 Ma
gametophyte-dominant life cycle, in which cells of the plant carry PreꞒ Ꞓ O S D C P T J K PgN
only a single set of genetic information. Mid-Ordovician[1] to present

It is estimated that there are about 9000 species of liverworts.[4]

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 2–20  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 "Hepaticae" from Ernst Haeckel's
which they occur. They are distributed globally in almost every Kunstformen der Natur, 1904
available habitat, most often in humid locations although there are
Scientific classification
desert and Arctic species as well. Some species can be a nuisance in
shady greenhouses or a weed in gardens.[5] Kingdom: Plantae
Clade: Embryophytes
Division: Marchantiophyta
Contents Stotler & Stotl.-Crand.,
Physical characteristics 1977[2] emend. 2000[3]
Description
Life cycle Classes and orders
Asexual reproduction
Haplomitriopsida
Ecology
Classification Haplomitriales
Relationship to other plants Treubiales
Internal classification
Marchantiopsida
Economic importance
Blasiales
Gallery
Neohodgsoniales
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See also Sphaerocarpales

References Lunulariales
External links
Marchantiales
Jungermanniopsida
Physical characteristics
Pelliales
Pallaviciniales
Description
Fossombroniales
Most liverworts are small, measuring from 2–20 millimetres Pleuroziales
(0.08–0.8 in) wide with individual plants less than 10 centimetres
Metzgeriales
(4  in) long,[6] so they are often overlooked. The most familiar
liverworts consist of a prostrate, flattened, ribbon-like or Porellales
branching structure called a thallus (plant body); these liverworts
Ptilidiales
are termed thallose liverworts. However, most liverworts produce
flattened stems with overlapping scales or leaves in two or more Jungermanniales
ranks, the middle rank is often conspicuously different from the
outer ranks; these are called leafy liverworts or scale liverworts.[7][8] (See the gallery below for
examples.)

Liverworts can most reliably be distinguished from the apparently

similar mosses by their single-celled rhizoids.[9] Other differences
are not universal for all mosses and all liverworts;[8] 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.[10][11] 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
A thallose liverwort, Lunularia cytoplasm of all other plants being unenclosed.[12] The overall
cruciata 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, like other bryophytes, have a gametophyte-dominant life cycle, with the sporophyte
dependent on the gametophyte.[12] 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 vascular 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.[13] 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.[14] Even in mosses, the sporophyte is more persistent and in hornworts, the sporophyte disperses
spores over an extended period.

Life cycle

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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.[15][16] 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). [8] 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. Sexual life cycle of a Marchantia-like liverwort
In dioicous liverworts, female and male sex organs are
borne on different and separate gametophyte plants. In
monoicous liverworts, the two kinds of reproductive structures are borne on different branches of the
same plant.[17] 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,[18] 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.[19]

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.[18] 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."[20] For example, in Riccia, when the
older parts of the forked thalli die, the younger tips become separate individuals.[20]

Some thallose liverworts such as Marchantia polymorpha and Lunularia cruciata produce small disc-
shaped gemmae in shallow cups.[21] Marchantia gemmae can be dispersed up to 120  cm by rain
splashing into the cups.[22] In Metzgeria, gemmae grow at thallus margins.[23] Marchantia polymorpha

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is a common weed in greenhouses, often covering the entire surface of containers;[24]:230 gemma
dispersal is the "primary mechanism by which liverwort spreads throughout a nursery or
greenhouse."[24]: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.[25] As with most groups of living
plants, they are most common (both in numbers and species) in moist tropical areas.[26] 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).[8][27] However, since this grouping makes the Bryophyta paraphyletic, the liverworts
are now usually given their own division.[28] The use of the division name Bryophyta sensu lato is still
found in the literature, but more frequently the name is used in a restricted sense to include only the
mosses.

Two hypotheses on the phylogeny 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.[29] Among the earliest fossils believed to be liverworts are compression fossils of
Pallaviciniites from the Upper Devonian of New York.[30] These fossils resemble modern species in the
Metzgeriales.[31] 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.[32] 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][33]

Internal classification

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Bryologists classify liverworts in the division Marchantiophyta. This divisional name is based on the
name of the most universally recognized liverwort genus Marchantia.[34] 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,[35] the Marchantiophyta may be subdivided into three classes:[36][37][38][39]

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).[36][40] It also includes the problematic genus Monoclea, which is sometimes placed in
its own order Monocleales.[41]
A third class, the Haplomitriopsida is newly recognized as the sister group of the other
liverworts;[39] it comprises the genera Haplomitrium, Treubia, and Apotreubia.

 
 
Haplomitriales
  Haplomitriopsida
   
 
 
Treubiales

 
  Blasiales
The diagram at right summarizes a
portion of a 2006 cladistic analysis Marchantiopsida  
Sphaerocarpales
   
of liverworts based upon three  
chloroplast genes, one nuclear Marchantiophyta  
   
gene, and one mitochondrial   Marchantiales
gene.[36]  
 
 
 
Metzgeriales (part)

Jungermanniopsida  
    Jungermanniales
 
 
 
 
Metzgeriales (part)

An updated classification by Söderström et al. 2016[42]

Marchantiophyta Stotler & Crandall-Stotler 2000

Haplomitriopsida Stotler & Crandall-Stotler 1977
Haplomitriales Hamlin 1972
Treubiales Schljakov 1972

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Marchantiopsida Cronquist, Takhtajan & Zimmermann 1966

Blasiidae He-Nygrén et al. 2006
Blasiales Stotler & Crandall-Stotler 2000
Marchantiidae Engler 1893 sensu He-Nygrén 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-Nygrén et al. 2006
Pelliales He-Nygrén 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 Klinggräff 1858

It is estimated that there are about 9000 species of liverworts, at least 85% of which belong to the leafy
group.[3][43] Despite that fact, no liverwort genomes have been sequenced to date and only few genes
identified and characterized.[44]

Economic importance
In ancient times, it was believed that liverworts cured diseases of the liver, hence the name.[45] In Old
English, the word liverwort literally means liver plant.[46] 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 hēpaticus 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".[47]

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
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A small collection of images showing liverwort structure and diversity:

Marchantia The archegonium of A sporophyte of Porella platyphylla

polymorpha, with Porella. Porella emerging clump growing on a
antheridial and from its archegonium. 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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(9th ed.). New York: W. H. Freeman. p. 599. ISBN 978-1429246446.
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External links
Liverwort structure in pictures (https://web.archive.org/web/20050323090944/http://www2.una.edu/pd
avis/hepatic_orders.htm)
LiToL: Assembling the Liverwort Tree of Life (https://web.archive.org/web/20070831024018/http://ww
w.biology.duke.edu/bryology/LiToL/) (note: for 500,000 million years ago read "480 million years
ago".)
Inter-relationships of Mosses, Liverworts, and Hornworts (https://web.archive.org/web/200504132144
51/http://www.bio.ilstu.edu/armstrong/syllabi/222book/chapt6.htm)
Additional information on Liverworts (https://web.archive.org/web/20051025142835/http://www.bio.u
mass.edu/biology/conn.river/liverwts.html)
Liverworts (https://web.archive.org/web/20050303043539/http://www.hcs.ohio-state.edu/hcs300/liver
1.htm)

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