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Marchantiales

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Marchantiales
Conocephalum conicum - a thallose liverwort
Scientific classification Edit this classification
Kingdom: Plantae
Division: Marchantiophyta
Class: Marchantiopsida
Subclass: Marchantiidae
Order: Marchantiales
Limpr., 1877[1]
Families

See Classification

Marchantiales is an order of thallose liverworts (also known as "complex thalloid 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 (a) special stalked vertical branches called archegoniophores or carpocephala, and (b) sterile cells called elaters inside the sporangium.[citation needed]

Phylogeny (extant Marchantiales)

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Based on the work by Villarreal et al. 2015[2]

Phylogeny (extant and extinct Marchantiales)

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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.[3]

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.[4] 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[5]). 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.[6] 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:[6]

Origin and evolution

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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.[3]

Molecular analyses, calibrated with the Triassic fossil Marchantites cyathodoides, suggest an origin for this group in the Permian [2] 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.[7] 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

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Taxonomy based on work by Söderström et al. 2016[8] and synonyms from Collection of genus-group names in a systematic arrangement.[9] The order Lunulariales, proposed by Long 2006,[10] has been recently re-included in Marchantiales as a family.[11][12]

Cross section through a marchantialian thallus.

References

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  1. ^ Limpricht, G. (1877). "Lebermoose". In Cohn, F. (ed.). Kryptogamen-Flora von Schlesien. Vol. 1. pp. 225–352.
  2. ^ a b Villarreal; et al. (2015). "Divergence times and the evolution of morphological complexity in an early land plant lineage (Marchantiopsida) with a slow molecular rate". New Phytologist. 209 (4): 1734–46. doi:10.1111/nph.13716. PMID 26505145.
  3. ^ a b Tomescu, Alexandru M.F.; Bomfleur, Benjamin; Bippus, Alexander C.; Savoretti, Adolfina (2018), "Why Are Bryophytes So Rare in the Fossil Record? A Spotlight on Taphonomy and Fossil Preservation", Transformative Paleobotany, Elsevier, pp. 375–416, doi:10.1016/b978-0-12-813012-4.00016-4, ISBN 978-0-12-813012-4, retrieved 2020-11-02
  4. ^ Anderson, Heidi (1976). "A review of the Bryophyta from the Upper Triassic Molteno Formation, Karroo Basin, South Africa". Palaeontologia Africana. 30: 21–30. hdl:10539/16189 – via WireDSpace.
  5. ^ Li, Ruiyun; Li, Xiaoqiang; Wang, Hongshan; Sun, Bainian (2019). "Ricciopsis sandaolingensis sp. nov., a new fossil bryophyte from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang, Northwest China". Palaeontologia Electronica. 22 (2). doi:10.26879/917.
  6. ^ a b Flores, Jorge R; Bippus, Alexander C; Suárez, Guillermo M; Hyvönen, Jaakko (2020). "Defying death: incorporating fossils into the phylogeny of the complex thalloid liverworts (Marchantiidae, Marchantiophyta) confirms high order clades but reveals discrepancies in family-level relationships". Cladistics. 16 (3): 231–247. doi:10.1111/cla.12442. PMID 34478198. S2CID 225165843.
  7. ^ Flores, Jorge R.; Bippus, Alexander C.; de Ullivarri, Carmen Fernández; Suárez, Guillermo M.; Hyvönen, Jaakko; Tomescu, Alexandru M. F. (December 2023). "Dating the evolution of the complex thalloid liverworts (Marchantiopsida): total-evidence dating analysis supports a Late Silurian-Early Devonian origin and post-Mesozoic morphological stasis". New Phytologist. 240 (5): 2137–2150. doi:10.1111/nph.19254. ISSN 0028-646X. PMID 37697646.
  8. ^ Söderström; et al. (2016). "World checklist of hornworts and liverworts". PhytoKeys (59): 1–826. doi:10.3897/phytokeys.59.6261. PMC 4758082. PMID 26929706.
  9. ^ "Part 2- Plantae (starting with Chlorophycota)". Collection of genus-group names in a systematic arrangement. Archived from the original on 6 October 2016. Retrieved 30 June 2016.
  10. ^ Long, D. G. (July 2006). "New Higher Taxa of Complex Thalloid Liverworts (Marchantiophyta – Marchantiopsida)". Edinburgh Journal of Botany. 63 (2–3): 257–262. doi:10.1017/S0960428606000606. ISSN 0960-4286.
  11. ^ Cole, Theodor C H; Hilger, Hartmut H; Goffinet, Bernard (24 May 2019). "Supplemental Information 1: Bryophyte Phylogeny Poster 2019 - full A0 size". doi:10.7287/peerj.preprints.27571v3/supp-1. {{cite journal}}: Cite journal requires |journal= (help)
  12. ^ Flores, Jorge R.; Catalano, Santiago A.; Muñoz, Jesus; Suárez, Guillermo M. (2018). "Combined phylogenetic analysis of the subclass Marchantiidae (Marchantiophyta): towards a robustly diagnosed classification". Cladistics. 34 (5): 517–541. doi:10.1111/cla.12225. hdl:10261/248464. ISSN 1096-0031. PMID 34706484. S2CID 52831959.
  • 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). ISBN 0-521-66097-1.
  • Grolle, Riclef (1983). "Nomina generica Hepaticarum; references, types and synonymies". Acta Botanica Fennica 121, 1-62.
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