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Marchantiales is an order of thallose
liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ...
s (also known as "complex thalloid liverworts") that includes species like ''
Marchantia polymorpha ''Marchantia polymorpha'' is a species of large thalloid liverwort in the class Marchantiopsida. ''M. polymorpha'' is highly variable in appearance and contains several subspecies. This species is dioicous, having separate male and female plants ...
'', a widespread plant often found beside rivers, and '' Lunularia cruciata'', a common and often troublesome weed in moist, temperate gardens and greenhouses. As in other
bryophyte The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
s, the
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
generation is dominant, with the sporophyte existing as a short-lived part of the
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring * Life-cycle hypothesis ...
, dependent upon the gametophyte. The
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''
Marchantia ''Marchantia'' is a genus of liverworts in the family Marchantiaceae and the order Marchantiales. The thallus of ''Marchantia'' shows differentiation into two layers: an upper photosynthetic layer with a well-defined upper epidermis with por ...
'' is often used to typify the order, although there are also many species of '' Asterella'' and species of the genus ''
Riccia ''Riccia'' is a genus of liverworts in the order Marchantiales. These plants are small and thalloid, that is not differentiated into root, stem and leaf. Depending on species, the thallus may be strap-shaped and about 0.5 to 4 mm wide with ...
'' are more numerous. The majority of genera are characterized by the presence of (a) special stalked vertical branches called archegoniophores or carpocephala, and (b) sterile cells celled
elater An elater is a cell (or structure attached to a cell) that is hygroscopic, and therefore will change shape in response to changes in moisture in the environment. Elaters come in a variety of forms, but are always associated with plant spores. In ...
s inside the sporangium.


Phylogeny (extant Marchantiales)

Based on the work by Villarreal et al. 2015


Phylogeny (extant and extinct Marchantiales)

Extinct complex thalloid liverworts are often represented by coalified compressions that preserve superficial morphological traits and do not allow exhaustively analysing their fine anatomy; though, in exceptional cases, fossils might preserve cell details. Generally speaking, extinct Marchantiales - which commonly date back to the Mesozoic - can be grouped in Marchantia-like and Riccia-like fossils according to their overall morphology. While the phylogenetic relationships among many extinct and extant Marchantiales remain equivocal, it has been suggested that some fossils are closely related to extant Marchantiales. Marchantites cyathodoides (Townrow) H. M. Anderson (Middle Triassic), for instance, is a Marchantia-like fossil whose detailed morphological characters (e.g., thallus with midrib, reduced air chambers, rhizoids and ventral scales) suggest a nested position within Marchantiales. Some Riccia-like fossils have even been assigned to families based on their overall morphology and branching patterns, such as the case of Ricciopsis sandaolingensis Li & Sun (Middle Jurassic). The first phylogenetic analyses that include both extinct and extant Marchantiales have further clarified the relationships among these taxa and have revealed new relationships among families. Likewise, the inclusion of fossils in total-evidence analyses implied that some groups of complex thalloid liverworts might be older than previously inferred. Summary tree based on the work by Flores et al. 2020:


Classification

Taxonomy based on work by Söderström et al. 2016 and synonyms from ''Collection of genus-group names in a systematic arrangement''. The order Lunulariales, proposed by Long 2006, has been recently re-included in Marchantiales as a family. *
Aytoniaceae Aytoniaceae is a family of liverworts in the order Marchantiales. Genera * '' Asterella'' Palisot De Beauvisage 1805 non Saccardo 1891 non Hara 1936 non Sollas 1886 * '' Cryptomitrium'' Austin ex Underwood 1884 * ''Mannia ''Mannia'' is a g ...
Cavers 1911 ebouliaceae; Grimaldiaceae** '' Asterella'' Palisot De Beauvisage 1805 'Fimbraria''_Nees_1820_non_''Fimbriaria_(flatworm).html" ;"title="Fimbraria.html" ;"title="'Fimbraria">'Fimbraria'' Nees 1820 non ''Fimbriaria (flatworm)">Fimbriaria'' Stackhouse 1809; '' Asterella'' section ''Wallichianae'' Long 2014; ''Hypenantron'' Corda 1829; ''Octokepos'' Griffith 1849] ** ''Cryptomitrium'' Austin ex Underwood 1884 [''Platycoaspis'' Lindberg 1889] ** '' Mannia'' Corda 1829 nom. cons. Cyathophora''_Gray_1821_non_Michelin_1843;_''Neesiella_(plant).html" ;"title="Cyathophora_(genus).html" ;"title="'Grimaldia'' Raddi 1818 non Schrank 1805; ''Cyathophora (genus)">Cyathophora'' Gray 1821 non Michelin 1843; ''Neesiella (plant)">Neesiella'' Schiffner 1893b; ''Duvalia'' Nees 1818 non Haworth 1812 non Bonpland 1813; ''Neesia'' Leman 1825 non Sprengel 1818; ''Arnelliella'' Massalongo 1914; ''Sindonisce'' Corda 1829] ** '' Plagiochasma (plant), Plagiochasma'' Lehmann & Lindenberg 1832 nom. cons. non Pomel 1883 'Aytonia''_Forster_&_Forster_1775;_'' 'Aytonia''_Forster_&_Forster_1775;_''Ruppina_(genus)">Ruppina''_Linnaeus;_''Rupinia.html" ;"title="Ruppina_(genus).html" ;"title="Aytonia.html" ;"title="'Aytonia">'Aytonia'' Forster & Forster 1775; ''Ruppina (genus)">Ruppina'' Linnaeus; ''Rupinia">Ruppina_(genus).html" ;"title="Aytonia.html" ;"title="'Aytonia">'Aytonia'' Forster & Forster 1775; ''Ruppina (genus)">Ruppina'' Linnaeus; ''Rupinia'' (sic) Corda 1829] ** ''Reboulia'' Raddi 1818 nom. cons. * Cleveaceae Cavers 1911 [Sauteriaceae] ** ''Athalamia'' Falconer 1848 ** ''Clevea'' Lindberg 1868 'Gollaniella''_Stephani_1905.html" ;"title="Gollaniella.html" ;"title="' 'Gollaniella''_Stephani_1905">Gollaniella.html"_;"title="'Gollaniella">'Gollaniella''_Stephani_1905**_''Peltolepis.html" ;"title="Gollaniella">'Gollaniella'' Stephani 1905">Gollaniella.html" ;"title="'Gollaniella">'Gollaniella'' Stephani 1905** ''Peltolepis">Gollaniella">'Gollaniella'' Stephani 1905">Gollaniella.html" ;"title="'Gollaniella">'Gollaniella'' Stephani 1905** ''Peltolepis'' Lindberg 1876 ** ''Sauteria'' Nees 1838 [''Sauchia'' Kashyap 1916] * Conocephalaceae Müller ex Grolle 1972 ** ''
Conocephalum ''Conocephalum'' is a genus of complex thalloid liverworts in the order Marchantiales and is the only extant genus in the family Conocephalaceae.AKIYAMA, H. (2022). Morphological and ecological diversification of Conocephalum conicum complex in Ja ...
'' Hill 1773 nom. cons. ['' Conicephala'' (sic) Wiggers 1780; '' Conocephalus'' (sic) Necker ex Dumortier 1822 non Blume 1825 non Thunberg 1815; '' Anthoconum'' Palisot De Beauvois 1804; '' Fegatella'' Raddi 1818; ''
Hepatica ''Hepatica'' (hepatica, liverleaf, or liverwort) is a genus of herbaceous perennials in the buttercup family, native to central and northern Europe, Asia and eastern North America. Some botanists include ''Hepatica'' within a wider interpre ...
'' Adanson 1763 non Miller 1754; '' Hepaticella'' Leman 1821; '' Strozzius'' Gray 1821; '' Nemoursia'' Merat 1840; '' Sandea'' Lindberg 1884; ''
Conocephalum ''Conocephalum'' is a genus of complex thalloid liverworts in the order Marchantiales and is the only extant genus in the family Conocephalaceae.AKIYAMA, H. (2022). Morphological and ecological diversification of Conocephalum conicum complex in Ja ...
'' ('' Sandea'') (Lindberg 1884) Inoue 1976] *
Corsiniaceae Corsiniaceae is a family of liverworts in the order Marchantiales Marchantiales is an order of thallose liverworts (also known as "complex thalloid liverworts") that includes species like ''Marchantia polymorpha'', a widespread plant often fou ...
Engler 1892 ** '' Corsinia'' Raddi 1818 Guentheria''_Leman_1821_non_Sprengel_1826;_''Tessellina.html" ;"title="Corsinia.html" ;"title="'Corsinia">Guentheria'' Leman 1821 non Sprengel 1826; ''Tessellina">Corsinia.html" ;"title="'Corsinia">Guentheria'' Leman 1821 non Sprengel 1826; ''Tessellina'' Dumortier 1822 non Dumortier 1874; ''Brissocarpus'' Lindenberg 1829] ** ''Cronisia'' Berkeley 1857 [''Carringtonia'' Lindberg 1868; ''Funicularia'' Trevisan 1877; ''Boschia'' Montagne 1856 non Korthals 1844; '' Myriorrhynchus'' Lindberg 1884] * Cyathodiaceae Stotler & Crandall-Stotler 2000 ** '' Cyathodium'' Kunze ex Lehmann 1834 'Synhymenium''_Griffith_1849;_''Synymenium.html" ;"title="Synhymenium.html" ;"title="'Synhymenium">'Synhymenium'' Griffith 1849; ''Synymenium">Synhymenium.html" ;"title="'Synhymenium">'Synhymenium'' Griffith 1849; ''Synymenium'' (sic) Hagen 1910; '' Cyathodium'' (''Metacyathodium'') Srivastava & Dixit 1996] * Dumortieraceae Long 2006 ** ''Dumortiera'' Nees 1824 * Exormothecaceae Müller ex Grolle 1972 ** '' Aitchisoniella'' Kashyap 1914 ** '' Exormotheca'' Mitten 1870 'Corbierella''_Douin_&_Trabut_1919.html" ;"title="Corbierella.html" ;"title="' 'Corbierella''_Douin_&_Trabut_1919">Corbierella.html"_;"title="'Corbierella">'Corbierella''_Douin_&_Trabut_1919**_''Stephensoniella_(plant).html" ;"title="Corbierella">'Corbierella'' Douin & Trabut 1919">Corbierella.html" ;"title="'Corbierella">'Corbierella'' Douin & Trabut 1919** ''Stephensoniella (plant)">Stephensoniella'' Kashyap 1914 non Cernosvitov 1934 non Lastochkin 1935 * Lunulariaceae Klingrräff 1858 **''Lunularia'' Adanson 1763 *Marchantiaceae Lindley 1836 ** ''Bucegia'' Radian 1903 [''Radiania'' Györffy 1927] ** ''
Marchantia ''Marchantia'' is a genus of liverworts in the family Marchantiaceae and the order Marchantiales. The thallus of ''Marchantia'' shows differentiation into two layers: an upper photosynthetic layer with a well-defined upper epidermis with por ...
'' Linnaeus 1753 'Chlamidium''_Corda_1828;_''Marchantiopsis.html" ;"title="Chlamidium.html" ;"title="'Chlamidium">'Chlamidium'' Corda 1828; ''Marchantiopsis">Chlamidium.html" ;"title="'Chlamidium">'Chlamidium'' Corda 1828; ''Marchantiopsis'' Gao & Chang 1982 non Douin & Douin 1918] ** ''Preissia'' Corda 1829 [''Achiton (genus), Achiton'' Corda 1829] * Monocleaceae Frank 1877 ** ''Monoclea'' Hooker 1820 * Monosoleniaceae Inoue 1966 ** '' Monosolenium'' Griffith 1849 'Dumortieropsis''_Horikawa_1934.html" ;"title="Dumortieropsis.html" ;"title="' 'Dumortieropsis''_Horikawa_1934">Dumortieropsis.html"_;"title="'Dumortieropsis">'Dumortieropsis''_Horikawa_1934 *_Oxymitraceae.html" ;"title="Dumortieropsis">'Dumortieropsis'' Horikawa 1934">Dumortieropsis.html" ;"title="'Dumortieropsis">'Dumortieropsis'' Horikawa 1934 * Oxymitraceae">Dumortieropsis">'Dumortieropsis'' Horikawa 1934">Dumortieropsis.html" ;"title="'Dumortieropsis">'Dumortieropsis'' Horikawa 1934 * Oxymitraceae Müller ex Grolle 1972 ** ''Oxymitra'' Bischoff ex Lindenberg 1829 non (Blume) Hooker & Thomson 1855 [''Pycnoscenus'' Lindberg 1863 ; ''Tessellina'' Dumortier 1874 non Dumortier 1822] * Ricciaceae Reichenbach 1828 ** ''
Riccia ''Riccia'' is a genus of liverworts in the order Marchantiales. These plants are small and thalloid, that is not differentiated into root, stem and leaf. Depending on species, the thallus may be strap-shaped and about 0.5 to 4 mm wide with ...
'' Linnaeus 1753 'Hemiseuma'' (Bischoff) von Klinggraeff 1858; ''
Riccia ''Riccia'' is a genus of liverworts in the order Marchantiales. These plants are small and thalloid, that is not differentiated into root, stem and leaf. Depending on species, the thallus may be strap-shaped and about 0.5 to 4 mm wide with ...
'' section ''Hemiseuma'' Bischoff; ''Hemiseumata'' Bischoff Lindley 1847; ''
Riccia ''Riccia'' is a genus of liverworts in the order Marchantiales. These plants are small and thalloid, that is not differentiated into root, stem and leaf. Depending on species, the thallus may be strap-shaped and about 0.5 to 4 mm wide with ...
'' ('' Pteroriccia'') (Schuster 1984) Schuster 1985; '' Pteroriccia'' Schuster 1984; '' Thallocarpus'' Lindberg 1874a; '' Cryptocarpus'' Austin 1869 non Kunth 1817 non Dozy & Molk. ex Dozy & Molk. 1846; '' Angiocarpus'' Trevisan 1877; '' Ricciella'' Braun 1821; '' Riccinia'' Trabut 1916] ** '' Ricciocarpos'' Corda 1829 'Euriccia''_Lacouture_1905;_''Lichenoides.html" ;"title="Euriccia.html" ;"title="'Euriccia">'Euriccia'' Lacouture 1905; ''Lichenoides">Euriccia.html" ;"title="'Euriccia">'Euriccia'' Lacouture 1905; ''Lichenoides'' Lindley 1847 non Hoffmann 1789 non Barrande 1846; ''Lemna'' Rafinesque 1817 non Linnaeus 1753] * Targioniaceae Dumortier 1829 ** ''Targionia (plant), Targionia'' Linnaeus 1753 non non Hesse 1923 * Wiesnerellaceae Inoue 1976 ** '' Wiesnerella'' Schiffner 1896


References

* Crandall-Stotler, Barbara J. & Stotler, Raymond E. "Morphology and classification of the Marchantiophyta". page 63 ''in'' A. Jonathan Shaw & Bernard Goffinet (Eds.), ''Bryophyte Biology''. (Cambridge: Cambridge University Press:2000). . * Grolle, Riclef (1983). "Nomina generica Hepaticarum; references, types and synonymies". Acta Botanica Fennica 121, 1-62.


External links


Life cycle



Additional photos of speciesMorphobank project
{{Taxonbar, from=Q139099 Liverwort orders