Liverwort

Liverworts are a group of non-vascular land plants forming the division Marchantiophyta (/mɑːrˌkæntiˈɒfətə, -oʊˈfaɪtə/ (listen)). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information. The division name was derived from the genus name Marchantia, named after his father by French botanist Jean Marchant.

It is estimated that there are about 9000 species of liverwort.[1] 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 distinguished from mosses in having unique complex oil bodies of high refractive index.

Liverworts are typically small, usually from 2 to 20 mm (0.079 to 0.787 in) wide with individual plants less than 10 cm (3.9 in) 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 habitat, most often in humid locations although there are desert and Arctic species as well. Some species can be a nuisance in shady greenhouses or a weed in gardens.[2]

## Contents

* 1 Physical characteristics
* 1.1 Description
* 1.2 Life cycle
* 1.2.1 Asexual reproduction
* 1.2.2 Symbiosis
* 2 Ecology
* 3 Classification
* 3.1 Evolution
* 3.2 Relationship to other plants
* 3.3 Internal classification
* 3.3.1 Extinct taxa
* 4 Economic importance
* 5 Gallery
* 6 See also
* 7 References
* 8 External links

## Physical characteristics

### Description

Most liverworts are small, measuring from 2 to 20 millimetres (0.08 to 0.8 in) wide with individual plants less than 10 centimetres (4 in) long,[3] so they are often overlooked. The most familiar liverworts consist of a prostrate, flattened, ribbon-like or branching structure called a thallus (plant body); these liverworts are termed thallose liverworts. However, most liverworts produce flattened stems with 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.[4][5] (See the gallery below for examples.)

A thallose liverwort, Lunularia cruciata

Liverworts can most reliably be distinguished from the apparently similar mosses by their single-celled rhizoids.[6] Other differences are not universal for all mosses and all liverworts;[5] 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,[7][8] as well as frequent dichotomous branching, all point to the plant being a liverwort. With a few exceptions, all liverworts undergo polyplastidic meiosis, in contrast to mosses and hornworts which have monoplastidic meiosis.[9] 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 plants being unenclosed.[10] 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, like other bryophytes, have a gametophyte-dominant life cycle, with the sporophyte dependent on the gametophyte.[10] The sporophyte of many liverworts are non-photosynthetic, but there are also several that are photosynthetic to various degrees.[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 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.[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] In mosses, the sporophyte is more persistent and in hornworts, the sporophyte disperses spores over an extended period.[14]

### Life cycle

Sexual life cycle of a Marchantia-like liverwort

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 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).[5] 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 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] The sporophyte lacks an apical meristem, an auxin-sensitive point of divergence with other land plants some time in the Late Silurian/Early Devonian.[20][21] 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."[22] For example, in Riccia, when the older parts of the forked thalli die, the younger tips become separate individuals.[22]

Some thallose liverworts such as Marchantia polymorpha and Lunularia cruciata produce small disc-shaped gemmae in shallow cups.[23] Marchantia gemmae can be dispersed up to 120 cm by rain splashing into the cups.[24] In Metzgeria, gemmae grow at thallus margins.[25] Marchantia polymorpha is a common weed in greenhouses, often covering the entire surface of containers;[26]: 230 gemma dispersal is the "primary mechanism by which liverwort spreads throughout a nursery or greenhouse."[26]: 231

#### Symbiosis

Thalloid liverworts typically harbor symbiotic glomeromycete fungi which have arbuscular (cilia-bearing) rootlets resembling those in vascular plants. Species in the Aneuraceae, however, associate with basidiomycete fungi belonging to the genus Tulasnella, while leafy liverworts typically harbor symbiotic basidiomycete fungi belonging to the genus Serendipita.[27]

## 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.[28] As with most groups of living plants, they are most common (both in numbers and species) in moist tropical areas.[29] Liverworts are more commonly found in moderate to deep shade, though desert species may tolerate direct sunlight and periods of total desiccation.

## Classification

### Evolution

Epiphytic thalloid liverworts evolved during the Triassic to the Cretaceous.[30]

### 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).[5][31] Somewhat more recently, the liverworts were given their own division (Marchantiophyta),[32] as bryophytes became considered to be paraphyletic. However, the most recent phylogenetic evidence indicates that liverworts are indeed likely part of a monophyletic clade ("Bryophyta sensu lato" or "Bryophyta Schimp.") alongside mosses and hornworts.[33][34][35] Hence, it has been suggested that the liverworts should be de-ranked to a class called Marchantiopsida.[36] In addition, there is strong phylogenetic evidence to suggest that liverworts and mosses form a monophyletic subclade named Setaphyta.[37][38][39]

'Monophyletic bryophytes' model  | 'Liverworts plus mosses–basal' model

|

| embryophytes |  |  |

vascular plants

|

bryophytes |  |  |

hornworts

|

setaphytes |  |  |

mosses

|

liverworts

|  | embryophytes  |  |  |  |  |

vascular plants

|

hornworts

setaphytes |  |  |

mosses

|

liverworts

Two of the most likely models for bryophyte evolution.[39]

An important conclusion from these phylogenies is that the ancestral stomata appear to have been lost in the liverwort lineage.[37][34] Among the earliest fossils believed to be liverworts are compression fossils of Pallaviciniites from the Upper Devonian of New York.[40] These fossils resemble modern species in the Metzgeriales.[41] 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.[42] 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.[43][44]

### 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.[45] 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 is not to be mistakenly associated with flowering plant genus Hepatica, of the buttercup family Ranunculaceae. In addition, the name Hepaticophyta is frequently misspelled in textbooks as Hepatophyta.

Although there is no consensus among bryologists as to the classification of liverworts above family rank,[46] the Marchantiophyta may be subdivided into three classes:[47][48][49]

* 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).[47][50] It also includes the problematic genus Monoclea, which is sometimes placed in its own order Monocleales.[51]
* A third class, the Haplomitriopsida is newly recognized as the sister group of the other liverworts;[49] it comprises the genera Haplomitrium, Treubia, and Apotreubia.

Forrest 2006[47] | Cole, Hilger & Goffinet 2021 [52]

|

| Marchantiophyta  |  |  |  | Haplomitriopsida |  |  |

Haplomitriales

|

Treubiales

|  | Marchantiopsida  |  |  |

Blasiales

|

|  |  |

Marchantiales

|

Sphaerocarpales

Jungermanniopsida  |  |  |

Metzgeriales (part)

|

|  |  |

Metzgeriales (part)

|

Jungermanniales

|  |  |  |  | Haplomitriopsida |  |  |

Haplomitriales

|

Treubiales

|  | Marchantiopsida |  | Blasiidae  |

Blasiales

|

Marchantiidae  |  |  |

Neohodgsoniales

|

|  |  |

Sphaerocarpales

|

|  |  |

Lunulariales

|

Marchantiales

Jungermanniopsida |  |  |  | Pelliidae  |  |  |

Pelliales

|

|  |  |

Fossombroniales

|

Pallaviciniales

|  | Metzgeriidae  |  |  |

Metzgeriales

|

Pleuroziales

Jungermanniidae  |  |  |

Jungermanniales

|

|  |  |

Porellales

|

Ptilidiales

An updated classification by Söderström et al. 2016[53]

* Marchantiophyta Stotler & Crandall-Stotler 2000
* Haplomitriopsida Stotler & Crandall-Stotler 1977
* Calobryales Hamlin 1972
* Treubiales Schljakov 1972
* 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
* Lunulariales Long 2006
* Marchantiales Limpricht 1877 (complex thalloids)
* Neohodgsoniales Long 2006
* Sphaerocarpales Cavers 1910 (bottle liverworts)
* Jungermanniopsida Stotler & Crandall-Stotler 1977
* Jungermanniidae Engler 1893 (leafy liverworts)
* Jungermanniales von Klinggräff 1858
* Porellales Schljakov 1972
* Ptilidiales Schljakov 1972
* Metzgeriidae Bartholomew-Began 1990
* Metzgeriales Chalaud 1930
* Pleuroziales Schljakov 1972
* Pelliidae He-Nygrén et al. 2006
* Fossombroniales Schljakov 1972
* Pallaviciniales Frey & Stech 2005
* Pelliales He-Nygrén et al. 2006

It is estimated that there are about 9000 species of liverworts, at least 85% of which belong to the leafy group.[54][55] Despite that fact, no liverwort genomes have been sequenced to date and only few genes identified and characterized.[56]

#### Extinct taxa

There are several known fossil genera from this group that are not assigned to any extant class:[57]

* †Discites Harris 1931
* †Eohepatica Heard and Jones 1931
* †Jungermanniopsis Howe and Hollick 1922
* †Jungermannites Göppert 1845
* †Schizolepidella Halle 1913
* †Thallomia Heard and Jones 1931

## Economic importance

In ancient times, it was assumed that liverworts cured diseases of the liver, hence the name.[58] In Old English, the word liverwort literally means liver plant.[59] 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".[60]

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 polymorpha, with antheridial and archegonial stalks

* The archegonium of Porella

* A sporophyte of Porella emerging from its archegonium

* Porella platyphylla clump growing on a tree

* Pellia epiphylla, growing on moist soil

* Plagiochila asplenioides, a leafy liverwort

* Riccia fluitans, an aquatic thallose liverwort

* Conocephalum conicum, a large thallose liverwort

## See also

* Bryophyte
* Embryophyte

## References

1. ↑ "Liverworts Homepage | UNB". https://www.unb.ca/research/liverworts/.
2. ↑ Schuster, Rudolf M. (1992). The Hepaticae and Anthocerotae of North America. VI. Chicago: Field Museum of Natural History. p. 19. ISBN 0-914868-21-7.
3. ↑ Schuster, Rudolf M. The Hepaticae and Anthocerotae of North America, vol. I, pp. 243–244. (New York: Columbia University Press, 1966)
4. ↑ Kashyap, Shiv Ram. Liverworts of the Western Himalayas and the Panjab Plain, vol. I, p. 1. (New Delhi: The Chronica Botanica, 1929)
5. ↑ 5.0 5.1 5.2 5.3 Schofield, W. B. Introduction to Bryology, pp. 135–140. (New York: Macmillan, 1985). ISBN 0-02-949660-8.
6. ↑ Nehira, Kunito. "Spore Germination, Protonemata Development and Sporeling Development", p. 347 in Rudolf M. Schuster (Ed.), New Manual of Bryology, volume I. (Nichinan, Miyazaki, Japan: The Hattori Botanical Laboratory, 1983). ISBN 49381633045 `{{isbn}}`: Check `isbn` value: length (help).
7. ↑ Allison, K. W. & John Child. The Liverworts of New Zealand, pp. 13–14. (Dunedin: University of Otago Press, 1975).
8. ↑ Conard, Henry S. and Paul L. Redfearn, Jr. How to Know the Mosses and Liverworts, revised ed., pp. 12–23. (Dubuque, Iowa: William C. Brown Co., 1979) ISBN 0-697-04768-7
9. ↑ Renzaglia, K. S.; Ashton, N. W.; Suh, D. Y. (2023). "Sporogenesis in Physcomitrium patens: Intergenerational collaboration and the development of the spore wall and aperture". Frontiers in Cell and Developmental Biology 11. doi:10.3389/fcell.2023.1165293. PMID 37123413.
10. ↑ 10.0 10.1 Harold C. Bold, C. J. Alexopoulos, and T. Delevoryas. Morphology of Plants and Fungi, 5th ed., p. 189. (New York: Harper-Collins, 1987). ISBN 0-06-040839-1.
11. ↑ Volume 1, Chapter 11-1: Photosynthesis: The Process
12. ↑ Fosket, Donald E. Plant Growth and Development: A Molecular Approach, p. 27. (San Diego: Academic Press, 1994). ISBN 0-12-262430-0.
13. ↑ Hicks, Marie L. Guide to the Liverworts of North Carolina, p. 10. (Durham: Duke University Press, 1992). ISBN 0-8223-1175-5.
14. ↑ Simpson, Michael G. (2019). "Evolution and Diversity of Green and Land Plants". Plant Systematics. pp. 55–74. doi:10.1016/b978-0-12-812628-8.50003-1. ISBN 978-0-12-812628-8.
15. ↑ Nehira, Kunito. "Spore Germination, Protonemata Development and Sporeling Development", pp. 358–374 in Rudolf M. Schuster (Ed.), New Manual of Bryology, volume I. (Nichinan, Miyazaki, Japan: The Hattori Botanical Laboratory, 1983). ISBN 49381633045 `{{isbn}}`: Check `isbn` value: length (help).
16. ↑ Chopra, R. N. & P. K. Kumra. Biology of Bryophytes, pp. 1–38. (New York: John Wiley & Sons, 1988). ISBN 0-470-21359-0.
17. ↑ Malcolm, Bill & Nancy Malcolm. Mosses and Other Bryophytes: An Illustrated Glossary, pp. 6 & 128\. (New Zealand: Micro-Optics Press, 2000). ISBN 0-473-06730-7.
18. ↑ 18.0 18.1 Campbell, Douglas H. The Structure and Development of Mosses and Ferns, pp. 73–74. (London: The Macmillan Co., 1918)
19. ↑ Pain, S. (2010). "Botanical ballistics". New Scientist 208 (2792/3): 45–47. doi:10.1016/s0262-4079(10)63177-6.
20. ↑ Cooke, Todd J.; Poli, Dorothybelle; Cohen, Jerry D. (2004). "Did auxin play a crucial role in the evolution of novel body plans during the Late Silurian-Early Devonian radiation of land plants?". The Evolution of Plant Physiology. pp. 85–107. doi:10.1016/b978-012339552-8/50006-8. ISBN 978-0-12-339552-8.
21. ↑ Friedman, William E.; Moore, Richard C.; Purugganan, Michael D. (2004). "The evolution of plant development". American Journal of Botany 91 (10): 1726–1741. doi:10.3732/ajb.91.10.1726. PMID 21652320. Bibcode: 2004AmJB...91.1726F.
22. ↑ 22.0 22.1 Lepp, Heino (15 April 2008). "Vegetative Reproduction". Australian Bryophytes. Australian National Botanic Gardens. http://www.anbg.gov.au/bryophyte/vegetative-reproduction.html.
23. ↑ Smith, AJE (1989) The Liverworts of Britain and Ireland, Cambridge University Press, Cambridge.
24. ↑ Equihua, C (1987). "Diseminacion de yemas en Marchantia polymorpha L. (Hepaticae)" (in fr). Cryptogamie, Bryologie, Lichénologie 8 (3): 199–217. INIST:8356670. https://www.biodiversitylibrary.org/page/60738107.
25. ↑ Lepp, Heino (28 February 2008). "Reproduction & Dispersal". Australian Bryophytes. Australian National Botanic Gardens. http://www.anbg.gov.au/bryophyte/sexual-vegetative.html.
26. ↑ 26.0 26.1 Newby, Adam; Altland, James E.; Gilliam, Charles H.; Wehtje, Glenn (December 2006). "Postemergence Liverwort Control in Container-Grown Nursery Crops". Journal of Environmental Horticulture 24 (4): 230–236. doi:10.24266/0738-2898-24.4.230.
27. ↑ Bidartondo, Martin I.; Duckett, Jeffrey G. (7 February 2010). "Conservative ecological and evolutionary patterns in liverwort–fungal symbioses". Proceedings of the Royal Society B: Biological Sciences 277 (1680): 485–492. doi:10.1098/rspb.2009.1458. PMID 19812075. Bibcode: 2010PBioS.277..485B.
28. ↑ Schuster, Rudolf M. The Hepaticae and Anthocerotae of North America, vol. I, pp. 243–249. (New York: Columbia University Press, 1966).
29. ↑ Pócs, Tamás. "Tropical Forest Bryophytes", p. 59 in A. J. E. Smith (Ed.) Bryophyte Ecology. (London: Chapman and Hall, 1982). ISBN 0-412-22340-6.
30. ↑ Bechteler, Julia; Schäfer-Verwimp, Alfons; Glenny, David; Cargill, D. Christine; Maul, Karola; Schütz, Nicole; von Konrat, Matt; Quandt, Dietmar et al. (December 2021). "The evolution and biogeographic history of epiphytic thalloid liverworts". Molecular Phylogenetics and Evolution 165. doi:10.1016/j.ympev.2021.107298. PMID 34464738. Bibcode: 2021MolPE.16507298B.
31. ↑ Crandall-Stotler, Barbara. & Stotler, Raymond E. "Morphology and classification of the Marchantiophyta". pp. 36–38 in A. Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology. (Cambridge: Cambridge University Press: 2000). ISBN 0-521-66097-1
32. ↑ Goffinet, Bernard. "Origin and phylogenetic relationships of bryophytes". pp. 124–149 in A. Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology. (Cambridge: Cambridge University Press:!2000). ISBN 0-521-66097-1
33. ↑ Leebens-Mack, James H. (2019). "One thousand plant transcriptomes and the phylogenomics of green plants". Nature 574 (7780): 679–685. doi:10.1038/s41586-019-1693-2. PMID 31645766.
34. ↑ 34.0 34.1 Harris, Brogan J. (2020). "Phylogenomic Evidence for the Monophyly of Bryophytes and the Reductive Evolution of Stomata". Current Biology 30 (11): P2201–2012.E2. doi:10.1016/j.cub.2020.03.048. PMID 32302587. Bibcode: 2020CBio...30E2001H.
35. ↑ Su, Danyan (2021). "Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants". Molecular Biology and Evolution 38 (8): 3332–3344. doi:10.1093/molbev/msab106. PMID 33871608.
36. ↑ de Sousa, Filipe (2019). "Nuclear protein phylogenies support the monophyly of the three bryophyte groups (Bryophyta Schimp.)". New Phytologist 222 (1): 565–575. doi:10.1111/nph.15587. PMID 30411803. Bibcode: 2019NewPh.222..565D.
37. ↑ 37.0 37.1 Puttick, Mark N. et al. (March 2018). "The Interrelationships of Land Plants and the Nature of the Ancestral Embryophyte". Current Biology 28 (5): 733–745.e2. doi:10.1016/j.cub.2018.01.063. PMID 29456145. Bibcode: 2018CBio...28E.733P.
38. ↑ Sousa, Filipe (2020). "The mitochondrial phylogeny of land plants shows support for Setaphyta under composition-heterogeneous substitution models". PeerJ 8 (4). doi:10.7717/peerj.8995. PMID 32377448.
39. ↑ 39.0 39.1 Cox, Cymon J. (2018). "Land Plant Molecular Phylogenetics: A Review with Comments on Evaluating Incongruence Among Phylogenies". Critical Reviews in Plant Sciences 37 (2–3): 113–127. doi:10.1080/07352689.2018.1482443. Bibcode: 2018CRvPS..37..113C.
40. ↑ Taylor, Thomas N. & Edith L. Taylor. The Biology and Evolution of Fossil Plants, p. 139. (Englewood Cliffs, NJ: Prentice Hall, 1993). ISBN 0-13-651589-4.
41. ↑ Oostendorp, Cora. The Bryophytes of the Palaeozoic and the Mesozoic, pp. 70–71. (Bryophytum Bibliotheca, Band 34, 1987). ISBN 3-443-62006-X.
42. ↑ VanAller Hernick, L.; Landing, E.; Bartowski, K. E. (2008). "Earth's oldest liverworts – Metzgeriothallus sharonae sp. nov. from the Middle Devonian (Givetian) of eastern New York, USA". Review of Palaeobotany and Palynology 148 (2–4): 154–162. doi:10.1016/j.revpalbo.2007.09.002. Bibcode: 2008RPaPa.148..154H.
43. ↑ Walker, Matt (October 12, 2010). "Fossils of earliest land plants discovered in Argentina" (in en-GB). BBC Earth. http://news.bbc.co.uk/earth/hi/earth_news/newsid_9079000/9079963.stm.
44. ↑ Rubinstein, C. V.; Gerrienne, P.; De La Puente, G. S.; Astini, R. A.; Steemans, P. (2010). "Early Middle Ordovician evidence for land plants in Argentina (eastern Gondwana)". New Phytologist 188 (2): 365–369. doi:10.1111/j.1469-8137.2010.03433.x. PMID 20731783. Bibcode: 2010NewPh.188..365R.
45. ↑ Crandall-Stotler, Barbara. & Stotler, Raymond E. "Morphology and classification of the Marchantiophyta". p. 63 in A. Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology. (Cambridge: Cambridge University Press:2000). ISBN 0-521-66097-1
46. ↑ Jones, E. W. (2004). Liverwort and Hornwort Flora of West Africa. Scripta Botnica Belgica. 30. Meise: National Botanic Garden (Belgium). p. 30. ISBN 90-72619-61-7.
47. ↑ 47.0 47.1 47.2 Forrest, Laura L.; Davis, E. Christine; Long, David G.; Crandall-Stotler, Barbara J.; Clark, Alexandra; Hollingsworth, Michelle L. (2006). "Unraveling the evolutionary history of the liverworts (Marchantiophyta): Multiple taxa, genomes and analyses". The Bryologist 109 (3): 303. doi:10.1639/0007-2745(2006)109[303:UTEHOT2.0.CO;2].
48. ↑ He-Nygrén, Xiaolan; Juslén, Aino; Ahonen, Inkeri; Glenny, David; Piippo, Sinikka (2006). "Illuminating the evolutionary history of liverworts (Marchantiophyta)—towards a natural classification". Cladistics 22 (1): 1–31. doi:10.1111/j.1096-0031.2006.00089.x. PMID 34892891.
49. ↑ 49.0 49.1 Renzaglia, Karen S.; Schuette, Scott; Duff, R. Joel; Ligrone, Roberto; Shaw, A. Jonathan; Mishler, Brent D.; Duckett, Jeffrey G. (2007). "Bryophyte phylogeny: Advancing the molecular and morphological frontiers". The Bryologist 110 (2): 179. doi:10.1639/0007-2745(2007)110[179:BPATMA2.0.CO;2].
50. ↑ Forrest, Laura L.; Barbara J. Crandall-Stotler (2004). "A Phylogeny of the Simple Thalloid Liverworts (Jungermanniopsida, Metzgeriidae) as Inferred from Five Chloroplast Genes". Monographs in Systematic Botany. Molecular Systematics of Bryophytes (Missouri Botanical Garden Press) 98: 119–140.
51. ↑ Schuster, Rudolf M. The Hepaticae and Anthocerotae of North America, vol. VI, p. 26. (Chicago: Field Museum of Natural History, 1992). ISBN 0-914868-21-7.
52. ↑ Cole, Theodor C. H.; Hilger, Hartmut H.; Goffinet, Bernard. "Bryophyte phylogeny poster: systematics and Characteristics of Nonvascular Land Plants (Mosses, Liverworts, Hornworts)". https://www.researchgate.net/publication/257240194.
53. ↑ Söderström (2016). "World checklist of hornworts and liverworts". PhytoKeys (59): 1–826. doi:10.3897/phytokeys.59.6261. PMID 26929706. Bibcode: 2016PhytK..59....1S.
54. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Crandall-Stotler-2000`
55. ↑ Sadava, David; David M. Hillis; H. Craig Heller; May Berenbaum (2009). Life: The Science of Biology (9th ed.). New York: W. H. Freeman. p. 599. ISBN 978-1429246446.
56. ↑ Sierocka, I; Kozlowski, L. P.; Bujnicki, J. M.; Jarmolowski, A; Szweykowska-Kulinska, Z (2014). "Female-specific gene expression in dioecious liverwort Pellia endiviifolia is developmentally regulated and connected to archegonia production". BMC Plant Biology 14 (1): 168. doi:10.1186/1471-2229-14-168. PMID 24939387. Bibcode: 2014BMCPB..14..168S.
57. ↑ "PBDB Taxon". https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=306729&is_real_user=1.
58. ↑ Dittmer, Howard J. Phylogeny and Form in the Plant Kingdom, p. 286. (Toronto: D. Van Nostrand Co., 1964)
59. ↑ Raven, P. H., R. F. Evert, & S. E. Eichhorn. Biology of Plants, 7th ed., p. 351. (New York: W. H. Freeman, 2005). ISBN 0-7167-1007-2.
60. ↑ Stern, Kingsley R. Introductory Plant Biology, 5th ed., p. 338. (Dubuque, Iowa: Wm. C. Brown Publishers, 1991) ISBN 0-697-09947-4.

## External links

* Data related to Marchantiophyta at Wikispecies
* Liverwort structure in pictures
* 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
* Additional information on Liverworts
* Liverworts

* v
* t
* e

Classification of Archaeplastida / Plantae sensu lato

Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta

Animal
Fungi)

Rhodophyta
(red algae)|

* Cyanidiophyceae
* Porphyridiophyceae
* Compsopogonophyceae
* Stylonematophyceae
* Rhodellophyceae
* Bangiophyceae
* Florideophyceae

Glaucocystophyta
(glaucophytes)|

* Glaucocystophyceae
* Glaucocystis
* Cyanophora
* Gloeochaete

Viridiplantae
(green algae,
& land plants)|

| M+C+S|

* Mesostigmatophyceae
* Chlorokybophyceae
* Spirotaenia

|

C+S|

| Chlorophyta
(prasinophytes)|

* Mamiellophyceae
* Nephrophyceae
* Palmophyllales
* Prasinophyceae
* Pseudoscourfieldiales
* Pyramimonadophyceae
* Scourfieldiales
* Pedinophyceae
* Chlorodendrophyceae
* UTC clade
* Ulvophyceae
* Trebouxiophyceae
* Chlorophyceae

|

Streptophyta
(charophytes,
& land plants)|

* Klebsormidiophyceae

Phragmo-
plastophyta|

* Charophyceae
* Coleochaetophyceae
* Zygnematophyceae
* Mesotaeniaceae

Embryophyta
(land plants)|

| Bryophytes
(non-vascular)|

* Marchantiophyta
* Anthocerotophyta
* Bryophyta "Moss"
* †Horneophytopsida

|

Tracheophyta
(vascular)|

| Lycopodiophyta
(microphylls)|

* †Zosterophyllopsida
* †Sawdoniales
* Isoetopsida
* Lycopodiopsida

|

Euphyllophyta
(megaphylls)|

| Polypodiopsida
(ferns)|

* †Cladoxylopsida
* †Stauropteridales
* †Zygopteridales
* Equisetidae
* Ophioglossidae
* Marattiidae
* Polypodiidae

|

Spermatophyta
(seed plants)|

* †Seed ferns
* Gymnosperms
* Gnetopsida
* Pinopsida
* Cycadopsida
* Ginkgoopsida
* Angiosperms or flowering plants
* Amborellales
* Nymphaeales
* Austrobaileyales
* Magnoliids
* Monocots
* Eudicots

Other|

* †Trimerophytopsida
* †Progymnosperm

Other|

* †Rhyniopsida

† = extinct. See also the list of plant orders.

Template:Marchantiophyta

* v
* t
* e

Extant life phyla/divisions by domain

| Bacteria|

* Acidobacteria
* Actinobacteria
* Aquificae
* Bacteroidetes
* Chlamydiae
* Chlorobi
* Chloroflexi
* Chrysiogenetes
* Cyanobacteria
* Deferribacteres
* Deinococcus–Thermus
* Dictyoglomi
* Fibrobacteres
* Firmicutes
* Fusobacteria
* Gemmatimonadetes
* Nitrospirae
* Planctomycetes
* Proteobacteria
* Spirochaetes
* Thermodesulfobacteria
* Thermomicrobia
* Thermotogae
* Verrucomicrobia

|

| Archaea|

* Crenarchaeota
* Euryarchaeota
* Korarchaeota
* Nanoarchaeota
* Archaeal Richmond Mine acidophilic nanoorganisms

|

Eukaryote|

| Protista|

* Heterokontophyta
* Haptophyta
* Cryptophyta
* Ciliophora
* Apicomplexa
* Dinoflagellata
* Euglenozoa
* Percolozoa
* Metamonada
* Radiolaria
* Foraminifera
* Cercozoa
* Rhodophyta
* Glaucophyta
* Amoebozoa

|

Fungi|

* Chytridiomycota
* Blastocladiomycota
* Neocallimastigomycota
* Glomeromycota
* Zygomycota
* Ascomycota
* Basidiomycota

Plant|

* Chlorophyta
* Charophyta
* Marchantiophyta
* Anthocerotophyta
* Moss
* Lycopodiophyta
* Pteridophyta
* Cycadophyta
* Ginkgophyta
* Pinophyta
* Gnetophyta
* Flowering plant

Animal|

* Sponge
* Ctenophora
* Placozoa
* Cnidaria
* Xenacoelomorpha
* Chordata
* Hemichordata
* Echinodermata
* Chaetognatha
* Kinorhyncha
* Loricifera
* Priapulida
* Nematoda
* Nematomorpha
* Onychophora
* Tardigrada
* Arthropoda
* Flatworm
* Gastrotricha
* Orthonectida
* Dicyemida
* Rotifera
* Acanthocephala
* Gnathostomulida
* Micrognathozoa
* Cycliophora
* Nemertea
* Phoronida
* Bryozoa
* Entoprocta
* Brachiopoda
* Mollusca
* Annelida

Incertae sedis|

Parakaryon

Wikidata ☰ Q189808 entry

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