Marchantiales | |
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Conocephalum conicum - a thallose liverwort | |
Scientific 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 ]
Based on the work by Villarreal et al. 2015 [2]
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]
Marchantiophyta |
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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 Marchantitesloreus 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.
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]
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 genetic information.
Marchantiopsida is a class of liverworts within the phylum Marchantiophyta. The species in this class are known as complex thalloid liverworts. The species in this class are widely distributed and can be found worldwide. Complex oil bodies are only found in the gametophyte.
Marchantiaceae is a family of liverworts in order Marchantiales. It contains a single genus Marchantia.
Lunularia is a genus of liverworts whose only species is Lunularia cruciata, the crescent-cup liverwort. Lunularia is either the only genus in the order Lunulariales, or may be placed in the order Marchantiales. The name, from Latin luna, moon, refers to the moon-shaped gemma cups.
Jungermanniales is the largest order of liverworts. They are distinctive among the liverworts for having thin leaf-like flaps on either side of the stem. Most other liverworts are thalloid, with no leaves. Due to their dorsiventral organization and scale-like, overlapping leaves, the Jungermanniales are sometimes called "scale-mosses".
Metzgeriales is an order of liverworts. The group is sometimes called the simple thalloid liverworts: "thalloid" because the members lack structures resembling stems or leaves, and "simple" because their tissues are thin and relatively undifferentiated. All species in the order have a small gametophyte stage and a smaller, relatively short-lived, spore-bearing stage. Although these plants are almost entirely restricted to regions with high humidity or readily available moisture, the group as a whole is widely distributed, and occurs on every continent except Antarctica.
Conocephalum is a genus of complex thalloid liverworts in the order Marchantiales and is the only extant genus in the family Conocephalaceae. Some species of Conocephalum are assigned to the Conocephalum conicum complex, which includes several cryptic species. Conocephalum species are large liverworts with distinct patterns on the upper thallus, giving the appearance of snakeskin. The species Conocephalum conicum is named for its cone-shaped reproductive structures, called archegoniophores. Common names include snakeskin liverwort, great scented liverwort and cat-tongue liverwort.
Jungermanniopsida is the largest of three classes within the division Marchantiophyta (liverworts).
Cavicularia densa is the only species in the liverwort genus Cavicularia. The species was first described in 1897 by Franz Stephani, and is endemic to Japan, where it grows on fine moist soil.
Pellia is a small but widespread genus of liverworts in the cool and temperate regions of the northern hemisphere. It is classified in order Pelliales and is a member of the family Pelliaceae within that order.
Haplomitriopsida is a class of liverworts comprising fifteen species in three genera. Recent cladistic analyses of nuclear, mitochondrial, and plastid gene sequences place this monophyletic group as the basal sister group to all other liverworts. The group thus provides a unique insight into the early evolution of liverworts in particular and of land plants in general.
Blasiales is an order of liverworts with a single living family and two species. The order has traditionally been classified among the Metzgeriales, but molecular cladistics suggests a placement at the base of the Marchantiopsida.
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. M. polymorpha has a wide distribution and is found worldwide. Common names include common liverwort or umbrella liverwort.
Aytoniaceae is a family of liverworts in the order Marchantiales.
Riella is a genus in the liverwort family Riellaceae, and includes about eighteen species. Plants in the genus are small and grow submerged in shallow temporary pools. Although the genus is widely distributed in the Northern Hemisphere, locating populations is often difficult. Its occurrence is sporadic and local, and the tiny plants are ephemeral. The ornamented spores remain viable for several years, allowing the plants to survive annual drying of their habitat. The plants are easily grown in laboratory cultures.
Pallaviciniales is an order of liverworts.
Ricciocarpos natans is the only species in the genus Ricciocarpos, a genus of liverworts in the family Ricciaceae. It was formerly listed in 1759 as a species of Riccia by Linnaeus, but then assigned to a new genus of its own in 1829 by August Carl Joseph Corda.
The oil bodies of liverworts, occasionally dubbed complex oil bodies for distinction, are unique organelles exclusive to the Marchantiophyta. They are markedly different from the oil bodies found in other land plants and algae in that they are membrane-bound, and not associated with food storage. The organelles are variable and present in an estimated 90% of liverwort species, often proving taxonomically relevant. As a whole, the formation and function of the organelles are poorly understood. Complex oil bodies are recognized as sites of isoprenoid biosynthesis and essential oil accumulation, and have been implicated with anti-herbivory, desiccation tolerance, and photo-protection.
Monoclea forsteri is one of the two species in the thallose liverwort family Monocleaceae. It is dioicous with the capsule dehiscing with a single longitudinal slit. Endemic and widely distributed throughout New Zealand, it is also the country's largest thalloid liverwort. Hooker described the species in 1820. The holotype is in the British Museum.
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