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Elizabeth A Brown, Royal Botanic Gardens Sydney, New South Wales, Australia
The liverworts are a group of green land plants that diverged before the dominant land plants such as angiosperms and gymnosperms. Together with mosses and hornworts, liverworts differ from all other land plants in having a free living gametophyte (a plant with only one set of chromosomes) as the dominant generation.
. Introduction . Families Included . Economically Important Species . Morphology . Ecology . Phylogeny
The liverworts are a group of green land plants that superﬁcially share a number of features with mosses, e.g. a life cycle with a dominant gametophyte generation, an apparently simple sporophyte (attached and substantially dependent on the gametophyte), a dependence on water for fertilization, small size, no secondary growth and rudimentary or no conductive tissues. Liverworts are found in most terrestrial and freshwater habitats, even quite extreme environments such as Antarctica. There are approximately 5000–8000 species in 300–350 genera and they are representatives of a group that were among the ﬁrst to colonize land some 400 plus million years ago. Although apparently simple, liverworts exhibit a range of morphologies and life strategies that allow them to successfully colonize and reproduce in a landscape dominated by the ﬂowering plants.
. Fossil History
signiﬁcant including Lejeuneaceae, Jubulaceae, Lepidoziaceae, Geocalycaceae, Schistochilaceae, Plagiochilaceae, Radulaceae, Jungermanniaceae s.l.). The Metzgeriales has about 30 genera and 13 families (Fossombroniaceae, Pelliaceae, Allisoniaceae, Phyllothalliaceae, Pallaviciniaceae, Sandeothallaceae, Makinoaceae, Blasiaceae, Aneuraceae, Vandiemeniaceae, Metzgeriaceae, Hymenophytaceae, Treubiaceae). The Treubiaceae is sometimes placed in an order of its own. The Calobryales (Haplomitriales) is a small group of c.10 species variously assigned to 1–3 genera (Haplomitrium, Calobryum, Steeriomitrium) in one family.
Economically Important Species
Bryophytes have an important role in the nitrogen, carbon and water cycles in many environments. From a human perspective, taxa such as Marchantia have been used medicinally in China, India and Europe. Other taxa such as Riccia have been used in the Himalayas and in Chinese herbal medicines. Liverworts have also been investigated chemically and have potential uses as pesticides and antibiotics against bacteria and fungi. Species of Frullania have been implicated in causing contact dermatitis in forestry workers.
The ranking and circumscription of some genera and higher level groupings vary in the classiﬁcation systems currently available. The Hepaticae (liverworts) are typically separated into two groups, the Jungermanniidae/ Jungermannopsida and the Marchantiidae/Marchantopsida (Table 1). The marchantioid group contains two orders, the Sphaerocarpales with two families (the Riellaceae and Sphaerocarpaceae with approximately 14 species in three genera) and the Marchantiales with about 250 species in 30 genera and 13 families (Cleveaceae, Aytoniaceae, Lunulariaceae, Conocephalaceae, Exormothecaceae, Marchantiaceae, Monoseleniaceae, Targioniaceae, Cyathodiaceae, Carrpaceae, Corsiniaceae, Oxymitraceae, Ricciaceae). The Monocleales, a small group containing one genus with two species, is variously placed in the marchantioid or jungermannioid lines on morphological evidence. Recent molecular work supports placement in the marchantioid group. The jungermannioids consist of three or four orders. About 85% of all liverworts belong to the Jungermanniales, which has some 200 genera and 40 families (the most
The liverworts have a life cycle similar to that of mosses and hornworts. The haploid gametophyte generation is the dominant free-living generation. The fundamental organization of the Hepaticae is fairly simple. There is an axis (sometimes ﬂattened into a thallus) growing from a single apical cell, with unicellular rhizoids, slime papillae (clubshaped cells that are sometimes elaborated) and male and female gametangia (which are usually restricted to particular zones). The spore germinates and produces a short-lived protonema that gives rise to a single gametophyte.
ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Peltolepis Family Aytoniaceae [Rebouliaceae] Genera Reboulia. Cryptomitrium Family Lunulariaceae Genus Lunularia Family Conocephalaceae Genera Conocephalum. Marchantia. Haplomitrium. Sewardiella Family Pelliaceae Genera Pellia. Sauteria. Ricciocarpus.els. Preissia. Geothallus Class Jungermanniopsida a Order Monocleales Family Monocleaceae Genus Monoclea Order Calobryales (Haplomitriales) Family Calobryaceae Genera Calobryum. Dumortiera. Greeneothallus Family Sandeothallaceae Genera Sandeothallus 2 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd. Asterella. Pteroriccia Order Sphaerocarpales Family Riellaceae Genus Riella Family Sphaerocarpaceae Genera Sphaercarpos. Podomitrium. Moerckia. Wiesnerella Family Exormothecaceae Genera Exormotheca.Bryophytes (Liverworts) Table 1 Classiﬁcation of the liverworts – phylum Marchantiophyta (Hepatophyta/Hepaticae) Class Marchantiopsida Order Marchantiales Family Cleveaceae [Sauteriaceae] Genera Athalamia. Hattorianthus. Noteroclada Family Allisoniaceae Genera Allisonia. Petalophyllum. Steeriomitrium Order Metzgeriales Family Fossombroniaceae Genera Fossombronia. Jensenia. Neohodgsonia.net . Plagiochasma. Corsinia Family Oxymitraceae Genus Oxymitra Family Ricciaceae Genera Riccia. ?Stephensoniella Family Marchantiaceae Genera Bucegia. Calycularia Family Phyllothalliaceae Genera Phyllothallia Family Pallaviciniaceae Genera Pallavicinia. Marchantiopsis Family Monoseleniaceae Genus Monoselenium Family Targioniaceae Genus Targionia Family Cyathodiaceae Genus Cyathodium Family Carrpaceae [Monocarpaceae] Genus Carrpos [Monocarpus] Family Corsiniaceae Genera Cronisia. Mannia. Nature Publishing Group / www.
Pedinophyllum. Tylimanthus. Platycaulis. Apotreubia Order Jungermanniales Family Herbertaceae Genera Herbertus. Isophyllaria. Lophonardia. Scapania. Herzogobryum.net 3 . Stephaniella. Diplocolea. Phragmatocolea. Accogynidium. Leiomitra. Gottschelia. Hepatostolonophora. Herzogiaria. Heteroscyphus. Cryptocolea.Bryophytes (Liverworts) Table 1 (Continued) Family Makinoaceae Genera Verdoornia. Geocalyx. Denotarisia. Chiastacaulon. Marsupella. Pseudolepicolea. Stenorrhipis. Chaetocolea. Lophozia. Nature Publishing Group / www. Gymnocoleopsis. Clasmatacolea. Notoscyphus. Leptoscyphopsis. Poeltia Family Scapaniaceae Genera Douinia. Krunodiplophyllum Family Delavayellaceae Genus Delavayella Family Geocalyaceae (c. Plagiochilidium. Conoscyphus. Archeochaete. Liochlaena. Cuspidatula. Sphenolobopsis. Gymnomitrion.els. Mylia Family Gymnomitriaceae Genera Acrolophozia. Evansianthus. Gongylanthus ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd. Eotrichocolea. Anomacaulis. Trichocolea. Marsupidium. Scaphophyllum. Arnellia. Cryptothallus Family Vandiemeniaceae Genus Vandiemenia Family Metzgeriaceae Genera Metzgeria. Roivainenia. Leptophyllopsis. Cryptocoleopsis. Gymnocolea. Pachyglossa. Anastrophyllum. Rhodoplagiochila Family Acrobolbaceae Genera Acrobolbus. Austrometzgeria. Lophochaete. Protosyzygiella. Pigafettoa. Anomylia. Xenochila. Harpanthus. Tritomaria. Xenocephalozia Family Plagiochilaceae Genera Plagiochila.22 genera) Genera Lophocolea. Cavicularia Family Aneuraceae Genera Aneura. Jamesoniella. Anastrepta. Makinoa Family Blasiaceae Genera Blasia. Saccogynidium. Jungermannia. Chiloscyphus. Leptoscyphus. Hattoria. Tetralophozia. Diplophyllum Family Blepharidophyllaceae Genera Blepharidophyllum. Goebelobryum Family Arnelliaceae Genera Southbya. Eremonotus. Lethocolea. Cryptochila. Austrolophozia. Plagiochilon. Gerhildiella. Andrewsianthus. Syzygiella. Tetracymbaliella. Triandrophyllum Family Trichocoleaceae Genera Temnoma. Riccardia. Archeophylla. Saccogyna. Acrochila. Mesoptychia. Prasanthus. Blepharostoma Family Grolleaceae Genus Grollea Family Trichotemnomaceae Genus Trichotemnoma Family Antheliaceae Genus Anthelia Family Vetaformaceae Genus Vetaforma Family Lepicoleaceae Genus Lepicolea Family Jungermanniaceae Genera Chandonanthus. Clandarium. Apometzgeria Family Hymenophytaceae Genus Hymenophytum b Family Treubiaceae Genera Treubia.
Micropterygium. Alobiellopsis. Pteropsiella. Kurzia. Cladopodiella. Trabacellula Family Cephaloziellaceae Genus Cephaloziella Family Jackiellaceae Genus Jackiella Family Adelanthaceae Genera Adelanthus. Cheilolejeunea. Stictolejeunea. Lejeunea. Neurolejeunea. Symbiezidium. Lopholejeunea. Neogrollea. Ascidiota. Amphijubula Family Lejeuneaceae Genera Bryopteris. Drepanolejeunea. Caudalejeunea. Leptolejeunea. Zoopsis. Archilejeunea. Paracromastigum. Chloranthelia.Bryophytes (Liverworts) Table 1 (Continued) Family Brevianthaceae Genus Brevianthus Family Chonecoleaceae Genus Chonecolea Family Schistochilaceae Genera Pleurocladopsis. Calyptrocolea. Odontoschisma. Mastigopelma. Isolembidium. Wettsteinia Family Ptilidiaceae Genus Ptilidium Family Mastigophoraceae Genus Mastigophora Family Chaetophyllopsidaceae Genera Chaetophyllopsis. Pachyschistochila Family Perssoniellaceae Genus Perssoniella Family Balantiopsidaceae Genera Ruizanthus. Frullania. Hygrolembidium. Paraschistochila.net . Acrolejeunea. Spruceanthus. Odontolejeunea. Ptychanthus. Isotachis. Dicranolejeunea. Schiﬀneriolejeunea. Thysananthus. Neesioscyphus. Trichocoleopsis Family Goebeliellaceae Genus Goebeliella Family Porellaceae Genera Porella. Taxilejeunea. Blepharolejeunea. Bazzania. Acromastigum. Megalembidium. Tuzibeanthus.els. Neohattoria. Drucella. Eoisotachis. Leucolejeunea. Ceratolejeunea. Metacalypogeia Family Cephaloziaceae (c. Marchesinia. Lepidolejeunea. Zoopsidella. Aureolejeunea. Phaeolejeunea. Sprucella. Pseudocephalozia. Anomoclada. Trachylejeunea. Lembidium. Macvicaria Family Jubulaceae Genera Jubula. Schiﬀneria. Harpalejeunea. Anisotachis Family Gyrothyraceae Genus Gyrothyra Family Lepidoziaceae Genera Lepidozia. Brachiolejeunea. Lepidogyna Family Jubulopsidaceae Genus Jubulopsis Family Neotrichocoleaceae Genera Neotrichocolea. Odontoseries. Iwaksukia. Hygrolejeunea. Verdoornianthus. Steerea. Amphilejeunea. Mytilopsis. Nature Publishing Group / www. Gackstroemia. 4 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd. Metahygrobiella. Herzogianthus Family Lepidolaenaceae Genera Lepidolaena. Schistochila.12 genera) Genera Cephalozia. Balantiopsis. Mastigolejeunea. Schusterella. Nowellia. ?Arachniopsis. Protocephalozia Family Phycolepidoziaceae Genus Phycolepidozia Family Calypogeiaceae Genera Calypogeia. Psiloclada. ?Hyalolepidozia. Rectolejeunea. Dendrobazzania. Omphalanthus. Telaranea. Acanthocoleus. Pycnolejeunea. Austrolembidium.
Siphonolejeunea. All of these features may be further elaborated by teeth or cilia (ﬁne hair-like structures.e. Schusterolejeunea. The spores are generally interspersed with elaters – long cells with spiral thickenings on the walls. coelocaules. Otolejeunea. The complex thalloids are internally diﬀerentiated into an upper (photosynthetic) layer of air chambers that open dorsally by an air pore and a lower parenchymatous region. Cryptolejeunea. but an array of transitional morphologies also occur. freeing the capsule from the protective gametophytic tissue. there is no columella. diﬀerent in shape and transversely inserted. Nature Publishing Group / www. Cyrtolejeunea. Myriocoleopsis. Colura. Eopleurozia a b Monocleales sometimes placed in the Marchantiopsida (see text). biﬁd (Herbertus) to many lobed (Lepidozia). Myriocolea. stem or receptacle-derived structures (perigynia. and traditionally has been separated into three types. Metzgeriopsis Family Radulaceae Genus Radula Family Pleuroziaceae Genera Pleurozia. Cephalantholejeunea. in the marchantioid line the walls are one cell thick and dehiscence is usually irregular). Cololejeunea. In Schistochila the leaves are keeled and bilobed while in Frullania the ventral lobe is formed into a sac that varies from helmet-shaped to elongate and pointed. The ventral surface has two or more rows of scales and two types of rhizoids (simple and peg). The antheridia are typically globose and stalked. The sterile jacket cells are not diﬀerentiated to form an apical cap (as occurs in moss dehiscence. Dactylophorella. Ophthalmolejeunea. (b) simple thallus and (c) complex thallus with air chambers. It is determinate in growth and is diﬀerentiated into a foot (the Riccia group excepted). The spiral thickening is hygroscopic and after capsule dehiscence twists the cell as it dries. Sphaerolejeunea. Cystolejeunea. Haplolejeunea.net . a seta (stalk) and a capsule. Physantholejeunea. Potamolejeunea. jungermannioid and marchantioid). The cells of the seta are thin-walled parenchyma and at spore maturity typically elongate up to 20 times their original length. In some liverworts the oil bodies are restricted to scattered oil cells. (a) leafy shoots. Echinocolea. These are membrane-bound organelles that are not found elsewhere in the plant kingdom and contain terpenoids. The capsule has a wall one to several cells thick and in most taxa dehiscence occurs by means of diﬀerentiated sutures (typically forming four valves. Placolejeunea. The lateral leaves may have a dorsal (Balantiopsis) or ventral (Radula) lobe or an inrolling of the leaf margin (Cheilolejeunea). Typically 5 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd. The sex organs are formed on the surface of the plant but may become enclosed in subsequent development. Cladiantholejeunea. Prionolejeunea. The two lateral rows of leaves may also be transversely inserted but most are inserted with a dorsal tilt (succubous) or a ventral tilt (incubous).g. Diplasiolejeunea.e. Calatholejeunea. Ecology Liverworts are constrained by a series of factors including the requirement of water for fertilization and the dependence of the sporophyte on the gametophyte. Macrolejeunea. Aphanotropis. There are three basic types of antheridial development (Haplomitrium/Calobryalian.Bryophytes (Liverworts) Table 1 (Continued) Rhaphidolejeunea. Nipponolejeunea. Liverworts lack various features found in mosses and/or hornworts. structures that occur around individual archegonia (pseudoperianths). Acantholejeunea. A vast array of morphologies that elaborate and confuse the underlying basic structure can be found in the leafy liverworts (Figure 1). Oil bodies are of some taxonomic interest but need to be observed in fresh material as they break down within a few hours of collecting. Crossotolejeunea. The archegonia are usually protected by some sort of enclosure that may consist of modiﬁed leaves (perianth). operculum or stomata (pores for gas exchange). Echinolejeunea. The gametophyte in the liverworts is considerably more malleable than in the mosses. Oil bodies occur in about 90% of liverwort taxa. Cladolejeunea. All the leaves may be similar but more usually the ventral row (known as underleaves or amphigastria) is smaller. Aphanolejeunea. Cyclolejeunea. Leiolejeunea. i. i. The leafy shoots consist of simple stems with two or three rows of leaves one cell thick. Stenolejeunea. they shed by rupturing of the jacket). The simple thallus varies from almost leafy (Fossombronia) to multistratose (many layered) and undiﬀerentiated (Aneura) or with a multistratose midrib and unistratose (single layered) wings (Metzgeria). Sometimes placed in an order of its own. Austrolejeunea. Trichocolea). Pictolejeunea. Nephelolejeunea. The leaves are arranged in two lateral rows with the third row ventral. assisting in the dispersal of the spores. marsupia) or a simple upgrowth of the thallus. Dactylolejeunea. In many genera of the marchantioids the archegonia are borne on erect specialized branches (archegoniophores). Amblyolejeunea. Cardiolejeunea. The sporophyte develops within the protection of the variously derived gametophytic tissues that also protect the archegonia (or are developed subsequent to fertilization). The leaves may be unlobed (Adelanthus). Tuymaella.els. e.
except for short periods in successional or ephemeral communities. club-shaped perianth protects the emerging sporophyte. tree bases and stream banks are obvious habitats but there are many more places where they can be found. Marchantia berteroana may be the dominant ground cover in early succession after a ﬁre. (f) Lateral leaf of Frullania with ventral lobule and stylus between lobule and stem.net . dorsal view. Nature Publishing Group / www. They may. the mosses.els. liverworts grow in moist sheltered microhabitats where there is less competition from the larger vascular plants. (i) and (j) Lepidozia species showing some of the variation in extent and number of lobes that may occur in liverworts. Phylogeny The evidence available suggests that the bryophytes consist of three groups. that have been evolutionarily separate from one another for a vast period of time. Moist shaded logs. particularly those that have a special life strategy. Many liverworts are capable of withstanding periods of desiccation or have developed an annual life cycle that allows them to persist as spores or subterranean tubers (Geothallus). ventral view showing underleaves with transverse insertion on the stem. (b) Incubous insertion. (c) Succubous leaf insertion.g. marginal budding of the thallus in Metzgeria 6 and regeneration from fragments (known in Herbertus but apparently rare in liverworts). (g) Lateral leaf of Cheilolejeunea with inrolled ventral lobule. most notably in wet habitats such as cloud forest at high elevations. This can occur by a variety of methods such as the decay of older regions of the plant with the younger regions becoming separated. form a conspicuous component. (e) Lateral leaf of Schistochila showing keel on lower edge of leaf and cilia on margin. (d) Radula. (h) Lateral leaf (half illustrated) of Trichocolea showing extensive lobing that occurs in some liverworts. Asexual methods of gametophyte reproduction appear to be extremely important in the development and maintenance of populations in many liverworts. Liverworts rarely dominate a habitat. ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd. Riccia. e. dorsal view. e.g. Such features result in taxa such as Riccia and Asterella being important components of soil crusts which prevent erosion in many arid environments. the splash cups of Marchantia. e. liverworts and hornworts. plants lack underleaves and lateral leaves have a ventral lobe.Bryophytes (Liverworts) Figure 1 (a) Incubous leaf insertion. The relationships of the liverworts are still unresolved with diﬀerent data sets giving somewhat diﬀerent answers. the production of a variety of specialized propagules (gemmae).g. however.
Radula.) New Manual of Bryology. Miocene amber deposits 20–30 million years old provide what many believe are the oldest records of living species. In: Shaw AJ and Goﬃnet B (eds) Bryophyte Biology. Kendrick P and Crane PR (1997) The Origin and Early Diversiﬁcation of Land Plants. Nichinan: The Hattori Botanical Laboratory. vols 1 and 2. ever more rigidly. Ann Arbor: The University of Michigan Herbarium.) New Manual of Bryology. Crum H (2001) Structural Diversity of Bryophytes. In the early Silurian there was a profound change. There has been no comprehensive phylogenetic analysis of the jungermannioids but interpretations such as those of Schuster are available. Further Reading Bischler H (1998) Systematics and evolution of the genera of the Marchantiales. Engelmann. e. Miller NG (1984) Tertiary and quaternary fossils. Solenostoma. provide controversial evidence that there was diversiﬁcation of embryophytes from the mid-Ordovician onwards. In: Bates JW. Pallaviciniites devonicus. one showing progressive morphological elaboration of structures in the gametophyte and the other a simpliﬁcation of the gametophyte (and sporophyte). In: Schuster RM (ed. Cambridge: Cambridge University Press. i. There are no other fossils known until the Carboniferous. Krassilov VA and Schuster RM (1984) Paleozoic and mesozoic fossils. that appeared in the record more than 350 million years ago. it is not until the early Tertiary (c. Endlicher S (1841) Enchiridion Botanicum. Treubiites and Metzgeriothallus). Naiaditaceae. The limited ultrastructural data on spore wall morphology is consistent with some of these spores having an aﬃnity with the liverworts. however. An Illustrated Glossary. In: Shaw AJ and Goﬃnet B (eds) Bryophyte Biology. In: Schuster RM (ed. which may have belonged to the Sphaerocarpales. Bryophytorum Bibliotheca 51: 1–129. Endlicher hypothesized that there has been a progressive elaboration of the sporophyte. Dispersed (unattached) spores. hornworts and other liverworts. Frullania. equivalent to the mosses. Within the liverworts there have been two alternative views of evolution. Malcolm B and Malcolm N (2000) Mosses and Other Bryophytes. A Cladistic Study. from forms such as the highly simpliﬁed capsule of Riccia to massive ones in the leafy liverworts. the neotropical Marchesinia bracteata and Stictolejeunea squamata. with some molecular data suggesting that the marchantioids are a separate lineage. Gottsche CM. Porella. Cambridge: Cambridge University Press. occur (Blasiites. In: Schuster RM (ed. Although there is an increase in the number of specimens that can be assigned to these groups during the Cretaceous.Bryophytes (Liverworts) Typically the liverworts and/or the hornworts are basal to the mosses and vascular plants or are resolved as sister groups to them. These simplistic views have also been challenged in recent works with the suggestion that there are two contrasting evolutionary histories in the marchantioids. Goﬃnet B (2000) Origin and phylogenetic relationships of bryophytes.e.g. Leeds: Maney and British Bryological Society. ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd. Nichinan: The Hattori Botanical Laboratory. The liverworts have been recognized traditionally as a single natural unit but even this view is being questioned. that the most derived groups are the most structurally simple ones. Fossil History There is little information on fossilized bryophytes available and to date the macrofossil record has contributed relatively little to the understanding of the transition from green algae to land plants. In: Shaw AJ and Goﬃnet B (eds) Bryophyte Biology. In: Schuster RM (ed. vols 1 and 2. of a series of malleable features. 50 million years ago) that species closely resembling modern leafy liverworts are recorded in Baltic amber (members of Cephalozia. The earliest known bryophyte is a late Devonian fossil.net 7 . Ashton NW and Duckett JG (eds) Bryology for the Twenty-ﬁrst Century. rather than being primitive. Nature Publishing Group / www. Micro-Optics Press: Nelson. Riccia.els. Nichinan: The Hattori Botanical Laboratory. with the ‘cryptospores’ being largely replaced by single trilete spores (consistent with a change from hepatic-like to early vascular plants or their progenitors). and one or two Jungermanniales. is more advanced. e. vols 1 and 2. Smithsonian Institution Press: Washington and London. development and classiﬁcation of hornworts. Bopp M and Capesius I (1998) A molecular approach to bryophyte systematics.) New Manual of Bryology. Schuster RM (1984) Comparative anatomy and morphology of the Hepaticae. Crandall-Stotler B and Stotler RE (2000) Morphology and classiﬁcation of the Marchantiophyta. in a number of groups ventral branching has been lost as part of an adaptation to tightly appressed growth on hard substrates along with the evolution of a complicatedly bilobed leaf that produces water pockets. Lindenberg JBG and Nees von Esenbeck CG (1844–1847) Synopsis Hepaticarum. Cambridge: Cambridge University Press. Leipzig: W. He suggests that one of the basic themes has been the ﬁxing. where three more taxa. Lepicolea. Marchantiales and an extinct family. Gottsche. Hamburg: Meissner.) New Manual of Bryology. There are records from the Permian that are attributed to the Marchantiales and subsequently a number of records from the Triassic – Metzgeriales. Scapania). It is known only from the gametophyte and has a form similar to that of the modern liverwort group Metzgeriales. Nichinan: The Hattori Botanical Laboratory. Renzaglia KS and Vaughn KC (2000) Anatomy. Schuster RM (1984) Evolution.g. vols 1 and 2. phylogeny and classiﬁcation of the Hepaticae. generally referred to the Metzgeriales. Lindenberg and Nees postulated the opposite.
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