Marchantiales


Marchantiales is an order of thallose liverworts that includes species like Marchantia polymorpha, 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 bryophytes, the gametophyte generation is dominant, with the sporophyte existing as a short-lived part of the life cycle, dependent upon the gametophyte.
The genus Marchantia is often used to typify the order, although there are also many species of Asterella and species of the genus Riccia are more numerous.
The majority of genera are characterized by the presence of special stalked vertical branches called archegoniophores or carpocephala, and sterile cells called elaters 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.
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 H. M. Anderson, for instance, is a Marchantia-like fossil whose detailed morphological characters 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. 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:

Origin and evolution

Liverworts, possess a compelling narrative of evolutionary intricacies, particularly within the realm of complex thalloid liverworts.
The genus Hepaticites, spanning Carboniferous strata of various regions, presents a puzzling case as its affiliation with the complex thalloid liverworts dependen on its individual species. Similarly, the Carboniferous Blasiites lobatus raises questions about its relationship with the Blasiales, the sister group to Marchantiales. The appearance of Marchantites ''loreus in the Early Permian of Russia offers the first clear evidence of the Marchantiales in the Paleozoic. However, rosette-shaped fossils that resemble Ricciaceae are as old as the Early Devonian – suggesting a much older origin for the group.
Molecular analyses, calibrated with the Triassic fossil Marchantites cyathodoides'', suggest an origin for this group in the Permian or later. In contrast, total-evidence dating paints a more ancient picture, tracing the complex thalloid liverworts back to the Silurian-Devonian boundary, highlighting a narrative of morphological stability across epochs. Thus, the complex thalloid liverworts emerge as significant players in the ongoing saga of plant evolution, their history intertwined with the deep complexities of geological time.

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 Cavers 1911
  • * Asterella Palisot De Beauvisage 1805
  • * Cryptomitrium Austin ex Underwood 1884
  • * Mannia Corda 1829 nom. cons.
  • * Plagiochasma Lehmann & Lindenberg 1832 nom. cons. non Pomel 1883
  • * Reboulia Raddi 1818 nom. cons.
  • Cleveaceae Cavers 1911
  • * Athalamia Falconer 1848
  • * Clevea Lindberg 1868
  • * Peltolepis Lindberg 1876
  • * Sauteria Nees 1838
  • Conocephalaceae Müller ex Grolle 1972
  • * Conocephalum Hill 1773 nom. cons.
  • Corsiniaceae Engler 1892
  • * Corsinia Raddi 1818
  • * Cronisia Berkeley 1857
  • Cyathodiaceae Stotler & Crandall-Stotler 2000
  • * Cyathodium Kunze ex Lehmann 1834
  • Dumortieraceae Long 2006
  • * Dumortiera Nees 1824
  • Exormothecaceae Müller ex Grolle 1972
  • * Aitchisoniella Kashyap 1914
  • * Exormotheca Mitten 1870
  • * Stephensoniella Kashyap 1914 non Cernosvitov 1934 non Lastochkin 1935
  • Lunulariaceae Klingrräff 1858
  • *Lunularia Adanson 1763
  • Marchantiaceae Lindley 1836
  • * Marchantia Linnaeus 1753
  • Monocleaceae Frank 1877
  • * Monoclea Hooker 1820
  • Monosoleniaceae Inoue 1966
  • * Monosolenium Griffith 1849
  • Oxymitraceae Müller ex Grolle 1972
  • * Oxymitra Bischoff ex Lindenberg 1829 non Hooker & Thomson 1855
  • Ricciaceae Reichenbach 1828
  • * Riccia Linnaeus 1753
  • * Ricciocarpos Corda 1829
  • Targioniaceae Dumortier 1829
  • * Targionia Linnaeus 1753 non non Hesse 1923
  • Wiesnerellaceae Inoue 1976
  • * Wiesnerella Schiffner 1896