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*Marchantia*, an example of a liverwort (Marchantiophyta)

Bryophytes (/ˈbraɪ.əˌfaɪts/)^[1] are a group of land plants (embryophytes), sometimes treated as a taxonomic division, that contains three groups of non-vascular land plants: the liverworts, hornworts, and mosses.^[2] In the strict sense, the division Bryophyta consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although some species can survive in drier environments.^[3] The bryophytes consist of about 20,000 plant species.^[4]^[5] Bryophytes produce enclosed reproductive structures (gametangia and sporangia), but they do not produce flowers or seeds. They reproduce sexually by spores and asexually by fragmentation or the production of gemmae.^[6]

Though bryophytes were considered a paraphyletic group in recent years, almost all of the most recent phylogenetic evidence supports the monophyly of this group, as originally classified by Wilhelm Schimper in 1879.^[7]

The term bryophyte comes from Ancient Greek βρύον (brúon) 'tree moss, liverwort' and φυτόν (phutón) 'plant'.

Features

The defining features of bryophytes are:

Their life cycles are dominated by a multicellular haploid gametophyte stage
Their sporophytes are diploid and unbranched
They do not have a true vascular tissue containing lignin (although some have specialized tissues for the transport of water)^[8]

Habitat

Bryophytes exist in a wide variety of habitats. They can be found growing in a range of temperatures (cold arctics and in hot deserts), elevations (sea-level to alpine), and moisture (dry deserts to wet rain forests). Bryophytes can grow where vascularized plants cannot because they do not depend on roots for uptake of nutrients from soil. Bryophytes can survive on rocks and bare soil.^[9]

Life cycle

Like all land plants (embryophytes), bryophytes have life cycles with alternation of generations.^[10] In each cycle, a haploid gametophyte, each of whose cells contains a fixed number of unpaired chromosomes, alternates with a diploid sporophyte, whose cells contain two sets of paired chromosomes. Gametophytes produce haploid sperm and eggs which fuse to form diploid zygotes that grow into sporophytes. Sporophytes produce haploid spores by meiosis, that grow into gametophytes.

Bryophytes are gametophyte dominant,^[11] meaning that the more prominent, longer-lived plant is the haploid gametophyte. The diploid sporophytes appear only occasionally and remain attached to and nutritionally dependent on the gametophyte.^[12] In bryophytes, the sporophytes are always unbranched and produce a single sporangium (spore producing capsule), but each gametophyte can give rise to several sporophytes at once.

Liverworts, mosses and hornworts spend most of their lives as gametophytes. Gametangia (gamete-producing organs), archegonia and antheridia, are produced on the gametophytes, sometimes at the tips of shoots, in the axils of leaves or hidden under thalli. Some bryophytes, such as the liverwort Marchantia, create elaborate structures to bear the gametangia that are called gametangiophores. Sperm are flagellated and must swim from the antheridia that produce them to archegonia which may be on a different plant. Arthropods can assist in transfer of sperm.^[13]

Fertilized eggs become zygotes, which develop into sporophyte embryos inside the archegonia. Mature sporophytes remain attached to the gametophyte. They consist of a stalk called a seta and a single sporangium or capsule. Inside the sporangium, haploid spores are produced by meiosis. These are dispersed, most commonly by wind, and if they land in a suitable environment can develop into a new gametophyte. Thus bryophytes disperse by a combination of swimming sperm and spores, in a manner similar to lycophytes, ferns and other cryptogams.

The sporophyte develops differently in the three groups. Both mosses and hornworts have a meristem zone where cell division occurs. In hornworts, the meristem starts at the base where the foot ends, and the division of cells pushes the sporophyte body upwards. In mosses, the meristem is located between the capsule and the top of the stalk (seta), and produces cells downward, elongating the stalk and elevating the capsule. In liverworts the meristem is absent and the elongation of the sporophyte is caused almost exclusively by cell expansion.^[14]

Sexuality

The arrangement of antheridia and archegonia on an individual bryophyte plant is usually constant within a species, although in some species it may depend on environmental conditions. The main division is between species in which the antheridia and archegonia occur on the same plant and those in which they occur on different plants. The term monoicous may be used where antheridia and archegonia occur on the same gametophyte and the term dioicous where they occur on different gametophytes.^[15]

In seed plants, "monoecious" is used where flowers with anthers (microsporangia) and flowers with ovules (megasporangia) occur on the same sporophyte and "dioecious" where they occur on different sporophytes. These terms occasionally may be used instead of "monoicous" and "dioicous" to describe bryophyte gametophytes. "Monoecious" and "monoicous" are both derived from the Greek for "one house", "dioecious" and "dioicous" from the Greek for two houses. The use of the "-oicy" terminology refers to the gametophyte sexuality of bryophytes as distinct from the sporophyte sexuality of seed plants.^[15]

Monoicous plants are necessarily hermaphroditic, meaning that the same plant produces gametes of both sexes.^[15] The exact arrangement of the antheridia and archegonia in monoicous plants varies. They may be borne on different shoots (autoicous), on the same shoot but not together in a common structure (paroicous or paroecious), or together in a common "inflorescence" (synoicous or synoecious).^[15]^[16] Dioicous plants are unisexual, meaning that an individual plant has only one sex.^[15] All four patterns (autoicous, paroicous, synoicous and dioicous) occur in species of the moss genus Bryum.^[16]

Classification and phylogeny

Traditionally, all living land plants without vascular tissues were classified in a single taxonomic group, often a division (or phylum). The term "Bryophyta" was first suggested by Braun in 1864.^[17] As early as 1879, the term Bryophyta was used by German bryologist Wilhelm Schimper to describe a group containing all three bryophyte clades (though at the time, hornworts were considered part of the liverworts).^[18]^[7] G.M. Smith placed this group between Algae and Pteridophyta.^[19] Although a 2005 study supported this traditional monophyletic view,^[20] by 2010 a broad consensus had emerged among systematists that bryophytes as a whole are not a natural group (i.e., are paraphyletic).^[21]^[22]^[23] However, a 2014 study concluded that these previous phylogenies, which were based on nucleic acid sequences, were subject to composition biases, and that, furthermore, phylogenies based on amino acid sequences suggested that the bryophytes are monophyletic after all.^[24] Since then, partially thanks to a proliferation of genomic and transcriptomic datasets, almost all phylogenetics studies based on nuclear and chloroplastic sequences have concluded that the bryophytes form a monophyletic group.^[24]^[25]^[18]^[26]^[27]^[28]^[29]^[30]^[31] Nevertheless, phylogenies based on mitochondrial sequences fail to support the monophyletic view.^[32]

The three bryophyte clades are the Marchantiophyta (liverworts), Bryophyta (mosses) and Anthocerotophyta (hornworts).^[33] However, it has been proposed that these clades are de-ranked to the classes Marchantiopsida, Bryopsida, and Anthocerotopsida, respectively.^[18] There is now strong evidence that the liverworts and mosses belong to a monophyletic clade, called Setaphyta.^[25]^[32]^[34]

Monophyletic view

The favored model, based on amino acids phylogenies, indicates bryophytes as a monophyletic group:^[24]

bryophytes

	mosses

	liverworts

Consistent with this view, compared to other living land plants, all three lineages lack vascular tissue containing lignin and branched sporophytes bearing multiple sporangia. The prominence of the gametophyte in the life cycle is also a shared feature of the three bryophyte lineages (extant vascular plants are all sporophyte dominant). However, if this phylogeny is correct, then the complex sporophyte of living vascular plants might have evolved independently of the simpler unbranched sporophyte present in bryophytes.^[24] Furthermore, this view implies that stomata evolved only once in plant evolution, before being subsequently lost in the liverworts.^[25]^[28]

Paraphyletic view

embryophytes

	vascular plants

	hornworts

setaphytes

	mosses

	liverworts

bryophytes

In this alternative view, the Setaphyta grouping is retained, but hornworts instead are sister to vascular plants.^[34] (Another paraphyletic view involves hornworts branching out first.)^[32]

Traditional morphology

Traditionally, when basing classifications on morphological characters, bryophytes have been distinguished by their lack of vascular structure. However, this distinction is problematic, firstly because some of the earliest-diverging (but now extinct) non-bryophytes, such as the horneophytes, did not have true vascular tissue, and secondly because many mosses have well-developed water-conducting vessels.^[35]^[36] A more useful distinction may lie in the structure of their sporophytes. In bryophytes, the sporophyte is a simple unbranched structure with a single spore-forming organ (sporangium), whereas in all other land plants, the polysporangiophytes, the sporophyte is branched and carries many sporangia.^[37]^[38] The contrast is shown in the cladogram below:^[39]

land plants

bryophytes

polysporangiophytes

	"protracheophytes", such as Aglaophyton or Horneophyton

	tracheophytes or vascular plants

Evolution

There have probably been several different terrestrialization events, in which originally aquatic organisms colonized the land, just within the lineage of the Viridiplantae.^[40] Between 510–630 million years ago, however, land plants emerged within the green algae.^[41] Molecular phylogenetic studies conclude that bryophytes are the earliest diverging lineages of the extant land plants.^[21]^[2]^[42]^[43] They provide insights into the migration of plants from aquatic environments to land. A number of physical features link bryophytes to both land plants and aquatic plants.^[44]

Similarities to algae and vascular plants

Green algae, bryophytes and vascular plants all have chlorophyll a and b, and the chloroplast structures are similar.^[45] Like green algae and land plants, bryophytes also produce starch stored in the plastids and contain cellulose in their walls.^[45] Distinct adaptations observed in bryophytes have allowed plants to colonize Earth's terrestrial environments. To prevent desiccation of plant tissues in a terrestrial environment, a waxy cuticle covering the soft tissue of the plant may be present, providing protection. In hornworts and mosses, stomata provide gas exchange between the atmosphere and an internal intercellular space system. The development of gametangia provided further protection specifically for gametes, the zygote and the developing sporophyte.^[46] The bryophytes and vascular plants (embryophytes) also have embryonic development which is not seen in green algae.^[45] While bryophytes have no truly vascularized tissue, they do have organs that are specialized for transport of water and other specific functions, analogous for example to the functions of leaves and stems in vascular land plants.^[45]

Bryophytes depend on water for reproduction and survival. In common with ferns and lycophytes, a thin layer of water is required on the surface of the plant to enable the movement of the flagellated sperm between gametophytes and the fertilization of an egg.^[46]

Comparative morphology

Summary of the morphological characteristics of the gametophytes of the three groups of bryophytes:

	Liverworts	Mosses	Hornworts
Structure	Thalloid or foliose	Foliose	Thalloid
Symmetry	Dorsiventral or radial	Radial	Dorsiventral
Rhizoids	Unicellular	Pluricellular	Unicellular
Chloroplasts/cell	Many	Many	One
Protonemata	Reduced	Present	Absent
Gametangia (antheridia and archegonia)	Superficial	Superficial	Immersed

Summary of the morphological characteristics of the sporophytes of the three groups of bryophytes:

	Liverworts	Mosses	Hornworts
Stomata	Absent	Present	Present
Structure	Small, without chlorophyll	Large, with chlorophyll	Large, with chlorophyll
Persistence	Ephemeral	Persistent	Persistent
Growth	Defined	Defined	Continuous
Apical growth^[47]	Absent	Present	Absent
Seta	Present	Present	Absent
Capsule form	Simple	Differentiated (operculum, peristome)	Elongated
Maturation of spores	Simultaneous	Simultaneous	Gradual
Dispersion of spores	Elaters	Peristome teeth	Pseudo-elaters
Columella	Absent	Present	Present
Dehiscence	Longitudinal or irregular	Transverse	Longitudinal

Uses

Environmental

Soil Conditioning
Bioindicators
Moss gardens
Pesticides

Characteristics of bryophytes make them useful to the environment. Depending on the specific plant texture, bryophytes have been shown to help improve the water retention and air space within soil.^[48] Bryophytes are used in pollution studies to indicate soil pollution (such as the presence of heavy metals), air pollution, and UV-B radiation.^[48] Gardens in Japan are designed with moss to create peaceful sanctuaries.^[48] Some bryophytes have been found to produce natural pesticides. The liverwort, Plagiochila, produces a chemical that is poisonous to mice.^[48] Other bryophytes produce chemicals that are antifeedants which protect them from being eaten by slugs.^[48] When Phythium sphagnum is sprinkled on the soil of germinating seeds, it inhibits growth of "damping off fungus" which would otherwise kill young seedlings.^[49]

Commercial

Fuel
Packaging
Wound Dressing

Peat is a fuel produced from dried bryophytes, typically Sphagnum. Bryophytes' antibiotic properties and ability to retain water make them a useful packaging material for vegetables, flowers, and bulbs.^[48] Also, because of its antibiotic properties, Sphagnum was used as a surgical dressing in World War I.^[48]

References

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Bibliography

Lesica, P.; McCune, B.; Cooper, S. V.; Hong, W. S. (1991). "Differences in lichen and bryophyte communities between old-growth and managed second-growth forests in the Swan Valley, Montana". Canadian Journal of Botany. 69 (8): 1745–1755. doi:10.1139/b91-222.

External links

Andrew's Moss Site Photos of bryophytes
27-May-2013 Centuries-old frozen plants revived, 400-year-old bryophyte specimens left behind by retreating glaciers in Canada are brought back to life in the laboratory.
Farge, Catherine La; Williams, Krista H.; England, John H. (22 May 2013). "Regeneration of Little Ice Age bryophytes emerging from a polar glacier with implications of totipotency in extreme environments". Proceedings of the National Academy of Sciences. 110 (24): 9839–9844. Bibcode:2013PNAS..110.9839L. doi:10.1073/pnas.1304199110. PMC 3683725. PMID 23716658.
Magill, R. E., ed. (1990). Glossarium polyglottum bryologiae. A multilingual glossary for bryology. Monographs in Systematic Botany from the Missouri Botanical Garden, v. 33, 297 pp. Online version: Internet Archive.

[1] "bryophyte". Dictionary.com Unabridged (Online). n.d.

[pmid12415314-2] Hedges, S. Blair (November 2002). "The origin and evolution of model organisms". Nature Reviews Genetics. 3 (11): 838–849. doi:10.1038/nrg929. PMID 12415314. S2CID 10956647.

[McMahon2012-3] Levetin, Estelle; McMahon, Karen (2012). Plants and Society. New York, NY: McGraw-Hill. p. 139. ISBN 978-0-07-352422-1.

[4] "Bryophytes (Mosses and liverworts) — The Plant List". theplantlist.org. Retrieved 2017-04-11.

[WhatrBryo-5] "What are Bryophytes". Southern Illinois University Carbondale. Archived from the original on 2018-10-19. Retrieved 2009-05-31.

[Vanderpoorten-6] Vanderpoorten, Alain; Goffinet, Bernard (2009). Introduction to Bryophytes. Cambridge: Cambridge University Press. p. 3. ISBN 978-0-511-54013-4.

[Schimp-7] Schimper, W.P. (1879). "Bryophyta". In Zittel, K.A. (ed.). Handbuch der Palaeontologie. Vol. 2. R. Oldenbourg.

[Lucasetal-8] Lucas, William J.; Groover, Andrew; Lichtenberger, Raffael; Furuta, Kaori; Yadav, Shri-Ram; Helariutta, Ykä; He, Xin-Qiang; Fukuda, Hiroo; Kang, Julie; Brady, Siobhan M.; Patrick, John W. (April 2013). "The Plant Vascular System: Evolution, Development and Functions F". Journal of Integrative Plant Biology. 55 (4): 294–388. doi:10.1111/jipb.12041. hdl:10261/76903. PMID 23462277.

[anbg-9] Lepp, Heino (28 February 2008). "Habitats". Australian Bryophytes. Australian National Botanic Gardens.

[Duckett-10] Ligrone, Roberto; Duckett, Jeffrey G.; Renzaglia, Karen S. (April 2012). "Major transitions in the evolution of early land plants: a bryological perspective". Annals of Botany. 109 (5): 851–871. doi:10.1093/aob/mcs017. PMC 3310499. PMID 22356739.

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@@ Line 3: / Line 3: @@
 [[File:Moss (Bryophyta) on the forest floor in Broken Bow, Oklahoma.jpg|thumb|An example of moss (Bryophyta) on the forest floor in Broken Bow, Oklahoma]]
-'''Bryophytes''' are a group of land plants, sometimes treated as a taxonomic [[Division (biology)|division]], that contains three groups of non-vascular land plants ([[embryophyte]]s): the [[Marchantiophyta|liverworts]], [[hornwort]]s and [[moss]]es.<ref name="pmid12415314">{{Cite journal |last=Hedges |first=S. Blair |date=November 2002 |title=The origin and evolution of model organisms |journal=Nature Reviews Genetics |volume=3 |issue=11 |pages=838–849 |doi=10.1038/nrg929 |pmid=12415314 |s2cid=10956647}}</ref> In the [[Sensu#Common_qualifiers|strict sense]], '''Bryophyta''' consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although they can survive in drier environments.<ref name="McMahon2012">{{Cite book |last=Levetin |first=Estelle |title=Plants and Society |last2=McMahon |first2=Karen |publisher=McGraw-Hill |year=2012 |isbn=978-0-07-352422-1 |location=New York, NY |page=139}}</ref> The bryophytes consist of about 20,000 plant species.<ref>{{Cite web |title=Bryophytes (Mosses and liverworts) — The Plant List |url=http://www.theplantlist.org/browse/B/ |access-date=2017-04-11 |website=www.theplantlist.org}}</ref><ref name="WhatrBryo">{{Cite web |title=What are Bryophytes |url=http://bryophytes.plant.siu.edu/bryojustified.html |url-status=dead |archive-url=https://web.archive.org/web/20181019163320/http://bryophytes.plant.siu.edu/bryojustified.html |archive-date=2018-10-19 |access-date=2009-05-31 |website=Southern Illinois University Carbondale}}</ref> Bryophytes produce enclosed reproductive structures (gametangia and sporangia), but they do not produce [[flower]]s or [[seed]]s. They reproduce sexually by [[spore]]s and asexually by fragmentation or the production of [[Gemma (botany)|gemmae]].<ref name="Vanderpoorten">{{Cite book |last=Vanderpoorten |first=Alain |title=Introduction to Bryophytes |last2=Goffinet |first2=Bernard |publisher=Cambridge University Press |year=2009 |isbn=978-0-511-54013-4 |location=Cambridge |page=3}}</ref> Though bryophytes were considered a [[paraphyletic]] group in recent years, almost all of the most recent [[Phylogenetics|phylogenetic]] evidence supports the [[monophyly]] of this group, as originally classified by [[Wilhelm Philippe Schimper|Wilhelm Schimper]] in 1879.<ref name="Schimp">{{Cite book |last=Schimper |first=W.P. |title=Handbuch der Palaeontologie |publisher=R. Oldenbourg |year=1879 |editor-last=Zittel |editor-first=K.A. |volume=2 |chapter=Bryophyta}}</ref> The term ''bryophyte'' comes {{etymology|grc|''[[wikt:βρύον|βρύον]]'' (brúon)|tree moss, liverwort||''[[wikt:φυτόν|φυτόν]]'' (phutón)|plant}}.
+'''Bryophytes''' ({{IPAc-en|'|b|r|aɪ|.|ə|ˌ|f|aɪ|t|s}})<ref>{{cite Dictionary.com|bryophyte}}</ref> are a group of [[embryophyte|land plants]] ([[embryophyte]]s), sometimes treated as a taxonomic [[division (biology)|division]], that contains three groups of [[non-vascular plant|non-vascular]] land plants: the [[Marchantiophyta|liverworts]], [[hornwort]]s, and [[moss]]es.<ref name="pmid12415314">{{Cite journal |last=Hedges|first=S. Blair |date=November 2002 |title=The origin and evolution of model organisms |journal=[[Nature Reviews Genetics]] |volume=3 |issue=11 |pages=838–849 |doi=10.1038/nrg929 |pmid=12415314 |s2cid=10956647}}</ref> In the [[Sensu#Common qualifiers|strict sense]], the division '''Bryophyta''' consists of the mosses only. Bryophytes are characteristically limited in size and prefer moist habitats although some species can survive in drier environments.<ref name="McMahon2012">{{Cite book |last1=Levetin |first1=Estelle |title=Plants and Society |last2=McMahon |first2=Karen |publisher=McGraw-Hill |year=2012 |isbn=978-0-07-352422-1 |location=New York, NY |page=139}}</ref> The bryophytes consist of about 20,000 plant species.<ref>{{Cite web |title=Bryophytes (Mosses and liverworts) — The Plant List |url=http://www.theplantlist.org/browse/B/ |access-date=2017-04-11 |website=theplantlist.org}}</ref><ref name="WhatrBryo">{{Cite web |title=What are Bryophytes |url=http://bryophytes.plant.siu.edu/bryojustified.html |url-status=dead |archive-url=https://web.archive.org/web/20181019163320/http://bryophytes.plant.siu.edu/bryojustified.html |archive-date=2018-10-19 |access-date=2009-05-31 |website=Southern Illinois University Carbondale}}</ref> Bryophytes produce enclosed reproductive structures ([[gametangia]] and [[sporangia]]), but they do not produce [[flower]]s or [[seed]]s. They reproduce sexually by [[spore]]s and asexually by fragmentation or the production of [[Gemma (botany)|gemmae]].<ref name="Vanderpoorten">{{Cite book |last1=Vanderpoorten |first1=Alain |title=Introduction to Bryophytes |last2=Goffinet |first2=Bernard |publisher=[[Cambridge University Press]] |year=2009 |isbn=978-0-511-54013-4 |location=Cambridge |page=3}}</ref>
+Though bryophytes were considered a [[paraphyletic]] group in recent years, almost all of the most recent [[phylogenetics|phylogenetic]] evidence supports the [[monophyly]] of this group, as originally classified by [[Wilhelm Philippe Schimper|Wilhelm Schimper]] in 1879.<ref name="Schimp">{{Cite book | last=Schimper | first=W.P. | title=Handbuch der Palaeontologie | publisher=R. Oldenbourg | year=1879 | editor-last=Zittel | editor-first=K.A. | volume=2 | chapter=Bryophyta}}</ref>
+The term ''bryophyte'' comes {{etymology|grc|''{{Wikt-lang|grc|βρύον}}'' ({{grc-transl|βρύον}})|tree moss, liverwort||''{{Wikt-lang|grc|φυτόν}}'' ({{grc-transl|φυτόν}})|plant}}.
 ==Features==
 The defining features of bryophytes are:
-* Their life cycles are dominated by a multicellular [[gametophyte]] stage
+* Their life cycles are dominated by a multicellular [[haploid]] [[gametophyte]] stage
-* Their [[sporophyte]]s are unbranched
+* Their [[sporophyte]]s are [[diploid]] and unbranched
-* They do not have a true [[vascular tissue]] containing [[lignin]] (although some have specialized tissues for the transport of water)<ref name="Lucasetal">{{Cite journal |last=Lucas |first=William J. |last2=Groover |first2=Andrew |last3=Lichtenberger |first3=Raffael |last4=Furuta |first4=Kaori |last5=Yadav |first5=Shri-Ram |last6=Helariutta |first6=Ykä |last7=He |first7=Xin-Qiang |last8=Fukuda |first8=Hiroo |last9=Kang |first9=Julie |last10=Brady |first10=Siobhan M. |last11=Patrick |first11=John W. |date=April 2013 |title=The Plant Vascular System: Evolution, Development and Functions F |journal=Journal of Integrative Plant Biology |volume=55 |issue=4 |pages=294–388 |doi=10.1111/jipb.12041 |pmid=23462277 |hdl-access=free |hdl=10261/76903}}</ref>
+* They do not have a true [[vascular tissue]] containing [[lignin]] (although some have specialized tissues for the transport of water)<ref name="Lucasetal">{{Cite journal |last1=Lucas |first1=William J. |last2=Groover |first2=Andrew |last3=Lichtenberger |first3=Raffael |last4=Furuta |first4=Kaori |last5=Yadav |first5=Shri-Ram |last6=Helariutta |first6=Ykä |last7=He |first7=Xin-Qiang |last8=Fukuda |first8=Hiroo |last9=Kang |first9=Julie |last10=Brady |first10=Siobhan M. |author-link10=Siobhan Brady |last11=Patrick |first11=John W. |date=April 2013 |title=The Plant Vascular System: Evolution, Development and Functions F |journal=Journal of Integrative Plant Biology |volume=55 |issue=4 |pages=294–388 |doi=10.1111/jipb.12041 |pmid=23462277 |hdl-access=free |hdl=10261/76903}}</ref>
 == Habitat ==
@@ Line 18: / Line 22: @@
 [[File:Moss alternation of generations 03-2012.png|thumb|550px|The life cycle of a dioicous bryophyte. The gametophyte (haploid) structures are shown in green, the sporophyte (diploid) in brown.]]
-Like all land plants (embryophytes), bryophytes have [[biological life cycle|life cycle]]s with [[alternation of generations]].<ref name="Duckett">{{Cite journal |last=Ligrone |first=Roberto |last2=Duckett |first2=Jeffrey G. |last3=Renzaglia |first3=Karen S. |date=April 2012 |title=Major transitions in the evolution of early land plants: a bryological perspective |journal=Annals of Botany |volume=109 |issue=5 |pages=851–871 |doi=10.1093/aob/mcs017 |pmc=3310499 |pmid=22356739}}</ref> In each cycle, a [[haploid]] [[gametophyte]], each of whose cells contains a fixed number of unpaired [[chromosome]]s, alternates with a [[diploid]] [[sporophyte]], whose cells contain two sets of paired chromosomes. Gametophytes produce haploid sperm and eggs which fuse to form diploid zygotes that grow into sporophytes. Sporophytes produce haploid spores by [[meiosis]], that grow into gametophytes.
+Like all land plants (embryophytes), bryophytes have [[biological life cycle|life cycle]]s with [[alternation of generations]].<ref name="Duckett">{{Cite journal |last1=Ligrone |first1=Roberto |last2=Duckett |first2=Jeffrey G. |last3=Renzaglia |first3=Karen S. |date=April 2012 |title=Major transitions in the evolution of early land plants: a bryological perspective |journal=Annals of Botany |volume=109 |issue=5 |pages=851–871 |doi=10.1093/aob/mcs017 |pmc=3310499 |pmid=22356739}}</ref> In each cycle, a haploid gametophyte, each of whose cells contains a fixed number of unpaired [[chromosome]]s, alternates with a diploid sporophyte, whose cells contain two sets of paired chromosomes. Gametophytes produce haploid sperm and eggs which fuse to form diploid zygotes that grow into sporophytes. Sporophytes produce haploid spores by [[meiosis]], that grow into gametophytes.
-Bryophytes are gametophyte dominant,<ref name="Carrington">{{Cite web |last=C.M. Sean Carrington |date=2013-11-04 |title=The Bryophytes |url=http://ecflora.cavehill.uwi.edu/bio_courses/bl14apl/bryo1.htm |publisher=The University of the West Indies}}</ref> meaning that the more prominent, longer-lived plant is the haploid gametophyte. The diploid sporophytes appear only occasionally and remain attached to and nutritionally dependent on the gametophyte.<ref>{{Cite journal |last=Johnson |first=M G |last2=Shaw |first2=A J |date=24 February 2016 |title=The effects of quantitative fecundity in the haploid stage on reproductive success and diploid fitness in the aquatic peat moss Sphagnum macrophyllum |journal=Heredity |volume=116 |issue=6 |pages=523–530 |doi=10.1038/hdy.2016.13 |pmc=4868265 |pmid=26905464}}</ref> In bryophytes, the sporophytes are always unbranched and produce a single [[sporangium]] (spore producing capsule), but each gametophyte can give rise to several sporophytes at once.
+Bryophytes are gametophyte dominant,<ref name="Carrington">{{Cite web |last=C.M. Sean Carrington |date=2013-11-04 |title=The Bryophytes |url=http://ecflora.cavehill.uwi.edu/bio_courses/bl14apl/bryo1.htm |publisher=The University of the West Indies |access-date=2020-03-05 |archive-date=2020-02-05 |archive-url=https://web.archive.org/web/20200205160203/http://ecflora.cavehill.uwi.edu/bio_courses/bl14apl/bryo1.htm |url-status=dead }}</ref> meaning that the more prominent, longer-lived plant is the haploid gametophyte. The diploid sporophytes appear only occasionally and remain attached to and nutritionally dependent on the gametophyte.<ref>{{Cite journal |last1=Johnson |first1=M G |last2=Shaw |first2=A J |date=24 February 2016 |title=The effects of quantitative fecundity in the haploid stage on reproductive success and diploid fitness in the aquatic peat moss Sphagnum macrophyllum |journal=Heredity |volume=116 |issue=6 |pages=523–530 |doi=10.1038/hdy.2016.13 |pmc=4868265 |pmid=26905464}}</ref> In bryophytes, the sporophytes are always unbranched and produce a single [[sporangium]] (spore producing capsule), but each gametophyte can give rise to several sporophytes at once.
-Liverworts, mosses and hornworts spend most of their lives as gametophytes. [[Gametangia]] (gamete-producing organs), [[Archegonium|archegonia]] and [[Antheridium|antheridia]], are produced on the gametophytes, sometimes at the tips of shoots, in the axils of leaves or hidden under thalli. Some bryophytes, such as the liverwort ''[[Marchantia]]'', create elaborate structures to bear the gametangia that are called gametangiophores. Sperm are flagellated and must swim from the antheridia that produce them to archegonia which may be on a different plant. Arthropods can assist in transfer of sperm.<ref name="Cronberg">{{Cite journal |last=Cronberg |first=N. |last2=Natcheva |first2=R. |last3=Hedlund |first3=K. |year=2006 |title=Microarthropods Mediate Sperm Transfer in Mosses |journal=Science |volume=313 |issue=5791 |page=1255 |doi=10.1126/science.1128707 |pmid=16946062 |s2cid=11555211}}</ref>
+Liverworts, mosses and hornworts spend most of their lives as gametophytes. [[Gametangia]] (gamete-producing organs), [[Archegonium|archegonia]] and [[Antheridium|antheridia]], are produced on the gametophytes, sometimes at the tips of shoots, in the axils of leaves or hidden under thalli. Some bryophytes, such as the liverwort ''[[Marchantia]]'', create elaborate structures to bear the gametangia that are called gametangiophores. Sperm are flagellated and must swim from the antheridia that produce them to archegonia which may be on a different plant. Arthropods can assist in transfer of sperm.<ref name="Cronberg">{{Cite journal |last1=Cronberg |first1=N. |last2=Natcheva |first2=R. |last3=Hedlund |first3=K. |year=2006 |title=Microarthropods Mediate Sperm Transfer in Mosses |journal=Science |volume=313 |issue=5791 |page=1255 |doi=10.1126/science.1128707 |pmid=16946062 |s2cid=11555211}}</ref>
 Fertilized eggs become zygotes, which develop into sporophyte embryos inside the archegonia. Mature sporophytes remain attached to the gametophyte. They consist of a stalk called a seta and a single sporangium or capsule. Inside the sporangium, haploid spores are produced by [[meiosis]]. These are dispersed, most commonly by wind, and if they land in a suitable environment can develop into a new gametophyte. Thus bryophytes disperse by a combination of swimming sperm and spores, in a manner similar to [[lycophyte]]s, [[fern]]s and other [[cryptogams]].
@@ Line 33: / Line 37: @@
 The arrangement of [[Antheridium|antheridia]] and [[Archegonium|archegonia]] on an individual bryophyte plant is usually constant within a species, although in some species it may depend on environmental conditions. The main division is between species in which the antheridia and archegonia occur on the same plant and those in which they occur on different plants. The term '''[[Monoicy|monoicous]]''' may be used where antheridia and archegonia occur on the same [[gametophyte]] and the term '''[[Dioicy|dioicous]]''' where they occur on different gametophytes.<ref name=GlimBisa14/>
-In [[seed plant]]s, "[[plant reproductive morphology|monoecious]]" is used where flowers with [[anther]]s (microsporangia) and flowers with [[ovule]]s (megasporangia) occur on the same [[sporophyte]] and "[[dioecy|dioecious]]" where they occur on different sporophytes. These terms occasionally may be used instead of "monoicous" and "dioicous" to describe bryophyte gametophytes. "Monoecious" and "monoicous" are both derived from the Greek for "one house", "dioecious" and "dioicous" from the Greek for two houses. The use of the "-oicy" terminology refers to the gametophyte sexuality of bryophytes as distinct from the sporophyte sexuality of seed plants.<ref name="GlimBisa14">{{Cite book |last=Glime |first=J.M. |url=http://www.bryoecol.mtu.edu |title=Bryophyte Ecology |last2=Bisang |first2=I. |date=2014 |publisher=Michigan Technological University and the International Association of Bryologists |editor-last=Glime |editor-first=J.M. |volume=1 Physiological Ecology |chapter=Sexuality: Its Determination (Ch. 3-1) |access-date=2014-11-09 |chapter-url=http://www.bryoecol.mtu.edu/chapters_VOL1/3-1Sexuality%20Its%20Determination.pdf |name-list-style=amp}}</ref>
+In [[seed plant]]s, "[[plant reproductive morphology|monoecious]]" is used where flowers with [[anther]]s (microsporangia) and flowers with [[ovule]]s (megasporangia) occur on the same [[sporophyte]] and "[[dioecy|dioecious]]" where they occur on different sporophytes. These terms occasionally may be used instead of "monoicous" and "dioicous" to describe bryophyte gametophytes. "Monoecious" and "monoicous" are both derived from the Greek for "one house", "dioecious" and "dioicous" from the Greek for two houses. The use of the "-oicy" terminology refers to the gametophyte sexuality of bryophytes as distinct from the sporophyte sexuality of seed plants.<ref name="GlimBisa14">{{Cite book |last1=Glime |first1=J.M. |url=http://www.bryoecol.mtu.edu |title=Bryophyte Ecology |last2=Bisang |first2=I. |date=2014 |publisher=Michigan Technological University and the International Association of Bryologists |editor-last=Glime |editor-first=J.M. |volume=1 Physiological Ecology |chapter=Sexuality: Its Determination (Ch. 3-1) |access-date=2014-11-09 |chapter-url=http://www.bryoecol.mtu.edu/chapters_VOL1/3-1Sexuality%20Its%20Determination.pdf |name-list-style=amp}}</ref>
-Monoicous plants are necessarily hermaphroditic, meaning that the same plant produces gametes of both sexes.<ref name=GlimBisa14/> The exact arrangement of the antheridia and archegonia in monoicous plants varies. They may be borne on different shoots (autoicous or [[Heteroecious|autoecious]]), on the same shoot but not together in a common structure (paroicous or paroecious), or together in a common "inflorescence" (synoicous or synoecious).<ref name=GlimBisa14/><ref name="Wats81">{{Cite book |last=Watson |first=E.V. |title=British Mosses and Liverworts |date=1981 |publisher=Cambridge University Press |isbn=9780521285360 |edition=3rd |page=7}} (Watson uses the "oecy" terms rather than the "oicy" terms.)</ref> Dioicous plants are '''unisexual''', meaning that an individual plant has only one sex.<ref name=GlimBisa14/> All four patterns (autoicous, paroicous, synoicous and dioicous) occur in species of the moss genus ''[[Bryum]]''.<ref name=Wats81/>
+Monoicous plants are necessarily hermaphroditic, meaning that the same plant produces gametes of both sexes.<ref name=GlimBisa14/> The exact arrangement of the antheridia and archegonia in monoicous plants varies. They may be borne on different shoots (autoicous), on the same shoot but not together in a common structure (paroicous or paroecious), or together in a common "inflorescence" (synoicous or synoecious).<ref name=GlimBisa14/><ref name="Wats81">{{Cite book |last=Watson |first=E.V. |title=British Mosses and Liverworts |date=1981 |publisher=Cambridge University Press |isbn=9780521285360 |edition=3rd |page=7}} (Watson uses the "oecy" terms rather than the "oicy" terms.)</ref> Dioicous plants are '''unisexual''', meaning that an individual plant has only one sex.<ref name=GlimBisa14/> All four patterns (autoicous, paroicous, synoicous and dioicous) occur in species of the moss genus ''[[Bryum]]''.<ref name=Wats81/>
 ==Classification and phylogeny==
-[[File:Hornwort (3144429129).jpg|thumb|Hornworts (Anthocrotophyta) were once believed to be the closest living relatives of the vascular plants.]]
+[[File:Hornwort (3144429129).jpg|thumb|Hornworts (Anthocerophyta) were once believed to be the closest living relatives of the vascular plants.]]
 [[File:Bryophyte sp Moss 3.jpg|thumb|Mosses are one group of bryophytes.]]
-Traditionally, all living land plants without vascular tissues were classified in a single taxonomic group, often a [[phylum|division]] (or phylum). The term "Bryophyta" was first suggested by Braun in 1864.<ref>{{Cite web |title=Phylum Bryophyta – Hierarchy |url=http://taxonomicon.taxonomy.nl/TaxonTree.aspx?src=0&id=3593 |publisher=Taxonomicon}}</ref> As early as 1879, the term Bryophyta was used by German bryologist Wilhelm Schimper to describe a group containing all three bryophyte clades (though at the time, hornworts were considered part of the liverworts).<ref name= Sousa2019/><ref name=Schimp/> [[Gilbert Morgan Smith|G.M. Smith]] placed this group between Algae and [[Pteridophyte|Pteridophyta]].<ref name="GMSmith">{{Cite book |last=Smith |first=G.M. |url=https://archive.org/details/cryptogamicbotan030182mbp |title=Cryptogamic Botany |date=1955 |publisher=McGraw-Hill |edition=2nd |volume=2 |location=New York}}</ref> Although a 2005 study supported this traditional monophyletic view,<ref>{{Cite journal |last=Goremykin |first=V. V. |last2=Hellwig |first2=F. H. |name-list-style=amp |year=2005 |title=Evidence for the most basal split in land plants dividing bryophyte and tracheophyte lineages |journal=Plant Systematics and Evolution |volume=254 |issue=1–2 |pages=93–103 |doi=10.1007/s00606-005-0337-1 |s2cid=41403901}}</ref> by 2010 a broad consensus had emerged among systematists that bryophytes as a whole are not a natural group (i.e., are [[paraphyletic]]).<ref name="Konrat">{{Cite journal |last=Konrat |first=M. |last2=Shaw |first2=A.J. |last3=Renzaglia |first3=K.S. |year=2010 |title=A special issue of ''Phytotaxa'' dedicated to Bryophytes: The closest living relatives of early land plants |journal=Phytotaxa |volume=9 |pages=5–10 |doi=10.11646/phytotaxa.9.1.3 |doi-access=free}}</ref><ref name="pmid18205621">{{Cite journal |last=Troitsky |first=A. V. |last2=Ignatov |first2=M. S. |last3=Bobrova |first3=V. K. |last4=Milyutina |first4=I. A. |date=December 2007 |title=Contribution of genosystematics to current concepts of phylogeny and classification of bryophytes |journal=Biochemistry (Moscow) |volume=72 |issue=12 |pages=1368–1376 |doi=10.1134/s0006297907120115 |pmid=18205621 |s2cid=13509400}}</ref><ref name="Knoop">{{Cite journal |last=Knoop |first=Volker |date=31 December 2010 |title=Looking for sense in the nonsense: a short review of non-coding organellar DNA elucidating the phylogeny of bryophytes |journal=Bryophyte Diversity and Evolution |volume=31 |issue=1 |pages=51–60 |doi=10.11646/bde.31.1.10}}</ref> However, a 2014 study concluded that these previous phylogenies, which were based on nucleic acid sequences, were subject to composition biases, and that, furthermore, phylogenies based on amino acid sequences suggested that the bryophytes are monophyletic after all.<ref name="CoxLiFostEmbl14" /> Since then, partially thanks to a proliferation of genomic and transcriptomic datasets, almost all phylogenetics studies based on nuclear and chloroplastic sequences have concluded that the bryophytes form a monophyletic group.<ref name="CoxLiFostEmbl14">{{Cite journal |last=Cox |first=Cymon J. |display-authors=etal |year=2014 |title=Conflicting Phylogenies for Early Land Plants are Caused by Composition Biases among Synonymous Substitutions |journal=Systematic Biology |volume=63 |issue=2 |pages=272–279 |doi=10.1093/sysbio/syt109 |pmc=3926305 |pmid=24399481}}</ref><ref name="Puttick2018">{{Cite journal |last=Puttick |first=Mark N. |display-authors=etal |year=2018 |title=The Interrelationships of Land Plants and the Nature of the Ancestral Embryophyte |journal=Current Biology |volume=28 |issue=5 |pages=733–745.e2 |doi=10.1016/j.cub.2018.01.063 |pmid=29456145 |hdl-access=free |s2cid=3269165 |hdl=1983/ad32d4da-6cb3-4ed6-add2-2415f81b46da}}</ref><ref name="Sousa2019">{{Cite journal |last=de Sousa |first=Filipe |display-authors=etal |year=2019 |title=Nuclear protein phylogenies support the monophyly of the three bryophyte groups (Bryophyta Schimp.) |journal=New Phytologist |volume=222 |issue=1 |pages=565–575 |doi=10.1111/nph.15587 |pmid=30411803 |hdl-access=free |hdl=1983/0b471d7e-ce54-4681-b791-1da305d9e53b |s2cid=53240320}}</ref><ref name="LeebensMack2019">{{Cite journal |last=Leebens-Mack |first=James H. |display-authors=etal |year=2019 |title=One thousand plant transcriptomes and the phylogenomics of green plants |journal=Nature |volume=574 |issue=7780 |pages=679–685 |doi=10.1038/s41586-019-1693-2 |pmc=6872490 |pmid=31645766}}</ref><ref name="Zhang2020">{{Cite journal |last=Zhang |first=Jian |display-authors=etal |year=2020 |title=The hornwort genome and early land plant evolution |journal=Nature Plants |volume=6 |issue=2 |pages=107–118 |doi=10.1038/s41477-019-0588-4 |pmc=7027989 |pmid=32042158}}</ref><ref name="Harris2020">{{Cite journal |last=Harris |first=Brogan J. |display-authors=etal |year=2020 |title=Phylogenomic Evidence for the Monophyly of Bryophytes and the Reductive Evolution of Stomata |journal=Current Biology |volume=30 |issue=11 |pages=P2201–2012.E2 |doi=10.1016/j.cub.2020.03.048 |pmid=32302587 |hdl-access=free |hdl=1983/fbf3f371-8085-4e76-9342-e3b326e69edd |s2cid=215798377}}</ref><ref name="Li2020">{{Cite journal |last=Li |first=Fay-Wei |display-authors=etal |year=2020 |title=Anthoceros genomes illuminate the origin of land plants and the unique biology of hornworts |journal=Nature Plants |volume=6 |issue=3 |pages=259–272 |doi=10.1038/s41477-020-0618-2 |pmc=8075897 |pmid=32170292 |hdl-access=free |hdl=10261/234303}}</ref><ref name="Sousa2020a">{{Cite journal |last=Sousa |first=Filipe |display-authors=etal |year=2020 |title=The Chloroplast Land Plant Phylogeny: Analyses Employing Better-Fitting Tree- and Site-Heterogeneous Composition Models |journal=Frontiers in Plant Science |volume=11 |page=1062 |doi=10.3389/fpls.2020.01062 |pmc=7373204 |pmid=32760416 |doi-access=free}}</ref><ref name="Su2021">{{Cite journal |last=Su |first=Danyan |display-authors=etal |year=2021 |title=Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants |journal=Molecular Biology and Evolution |volume=38 |issue=8 |pages=3332–3344 |doi=10.1093/molbev/msab106 |pmc=8321542 |pmid=33871608}}</ref> Nevertheless, phylogenies based on mitochondrial sequences fail to support the monophyletic view.<ref name="Sousa2020b">{{Cite journal |last=Sousa |first=Filipe |display-authors=etal |year=2020 |title=The mitochondrial phylogeny of land plants shows support for Setaphyta under composition-heterogeneous substitution models |journal=PeerJ |volume=8 |issue=4 |page=e8995 |doi=10.7717/peerj.8995 |pmc=7194085 |pmid=32377448}}</ref>
+Traditionally, all living land plants without vascular tissues were classified in a single taxonomic group, often a [[phylum|division]] (or phylum). The term "Bryophyta" was first suggested by Braun in 1864.<ref>{{Cite web |title=Phylum Bryophyta – Hierarchy |url=http://taxonomicon.taxonomy.nl/TaxonTree.aspx?src=0&id=3593 |publisher=Taxonomicon}}</ref> As early as 1879, the term Bryophyta was used by German bryologist Wilhelm Schimper to describe a group containing all three bryophyte clades (though at the time, hornworts were considered part of the liverworts).<ref name= Sousa2019/><ref name=Schimp/> [[Gilbert Morgan Smith|G.M. Smith]] placed this group between Algae and [[Pteridophyte|Pteridophyta]].<ref name="GMSmith">{{Cite book |last=Smith |first=G.M. |url=https://archive.org/details/cryptogamicbotan030182mbp |title=Cryptogamic Botany |date=1955 |publisher=McGraw-Hill |edition=2nd |volume=2 |location=New York}}</ref> Although a 2005 study supported this traditional monophyletic view,<ref>{{Cite journal |last1=Goremykin |first1=V. V. |last2=Hellwig |first2=F. H. |name-list-style=amp |year=2005 |title=Evidence for the most basal split in land plants dividing bryophyte and tracheophyte lineages |journal=Plant Systematics and Evolution |volume=254 |issue=1–2 |pages=93–103 |doi=10.1007/s00606-005-0337-1 |s2cid=41403901}}</ref> by 2010 a broad consensus had emerged among systematists that bryophytes as a whole are not a natural group (i.e., are [[paraphyletic]]).<ref name="Konrat2">{{Cite journal |last1=Konrat |first1=M. |last2=Shaw |first2=A.J. |last3=Renzaglia |first3=K.S. |year=2010 |title=A special issue of ''Phytotaxa'' dedicated to Bryophytes: The closest living relatives of early land plants |journal=Phytotaxa |volume=9 |pages=5–10 |doi=10.11646/phytotaxa.9.1.3 |doi-access=free}}</ref><ref name="pmid18205621">{{Cite journal |last1=Troitsky |first1=A. V. |last2=Ignatov |first2=M. S. |last3=Bobrova |first3=V. K. |last4=Milyutina |first4=I. A. |date=December 2007 |title=Contribution of genosystematics to current concepts of phylogeny and classification of bryophytes |journal=Biochemistry (Moscow) |volume=72 |issue=12 |pages=1368–1376 |doi=10.1134/s0006297907120115 |pmid=18205621 |s2cid=13509400}}</ref><ref name="Knoop">{{Cite journal |last=Knoop |first=Volker |date=31 December 2010 |title=Looking for sense in the nonsense: a short review of non-coding organellar DNA elucidating the phylogeny of bryophytes |journal=Bryophyte Diversity and Evolution |volume=31 |issue=1 |pages=51–60 |doi=10.11646/bde.31.1.10}}</ref> However, a 2014 study concluded that these previous phylogenies, which were based on nucleic acid sequences, were subject to composition biases, and that, furthermore, phylogenies based on amino acid sequences suggested that the bryophytes are monophyletic after all.<ref name="CoxLiFostEmbl14" /> Since then, partially thanks to a proliferation of genomic and transcriptomic datasets, almost all phylogenetics studies based on nuclear and chloroplastic sequences have concluded that the bryophytes form a monophyletic group.<ref name="CoxLiFostEmbl14">{{Cite journal |last=Cox |first=Cymon J. |display-authors=etal |year=2014 |title=Conflicting Phylogenies for Early Land Plants are Caused by Composition Biases among Synonymous Substitutions |journal=Systematic Biology |volume=63 |issue=2 |pages=272–279 |doi=10.1093/sysbio/syt109 |pmc=3926305 |pmid=24399481}}</ref><ref name="Puttick2018">{{Cite journal |last=Puttick |first=Mark N. |display-authors=etal |year=2018 |title=The Interrelationships of Land Plants and the Nature of the Ancestral Embryophyte |journal=Current Biology |volume=28 |issue=5 |pages=733–745.e2 |doi=10.1016/j.cub.2018.01.063 |pmid=29456145 |hdl-access=free |s2cid=3269165 |hdl=1983/ad32d4da-6cb3-4ed6-add2-2415f81b46da}}</ref><ref name="Sousa2019">{{Cite journal |last=de Sousa |first=Filipe |display-authors=etal |year=2019 |title=Nuclear protein phylogenies support the monophyly of the three bryophyte groups (Bryophyta Schimp.) |journal=New Phytologist |volume=222 |issue=1 |pages=565–575 |doi=10.1111/nph.15587 |pmid=30411803 |hdl-access=free |hdl=1983/0b471d7e-ce54-4681-b791-1da305d9e53b |s2cid=53240320}}</ref><ref name="LeebensMack2019">{{Cite journal |last=Leebens-Mack |first=James H. |display-authors=etal |year=2019 |title=One thousand plant transcriptomes and the phylogenomics of green plants |journal=Nature |volume=574 |issue=7780 |pages=679–685 |doi=10.1038/s41586-019-1693-2 |pmc=6872490 |pmid=31645766}}</ref><ref name="Zhang2020">{{Cite journal |last=Zhang |first=Jian |display-authors=etal |year=2020 |title=The hornwort genome and early land plant evolution |journal=Nature Plants |volume=6 |issue=2 |pages=107–118 |doi=10.1038/s41477-019-0588-4 |pmc=7027989 |pmid=32042158}}</ref><ref name="Harris2020">{{Cite journal |last=Harris |first=Brogan J. |display-authors=etal |year=2020 |title=Phylogenomic Evidence for the Monophyly of Bryophytes and the Reductive Evolution of Stomata |journal=Current Biology |volume=30 |issue=11 |pages=P2201–2012.E2 |doi=10.1016/j.cub.2020.03.048 |pmid=32302587 |hdl-access=free |hdl=1983/fbf3f371-8085-4e76-9342-e3b326e69edd |s2cid=215798377}}</ref><ref name="Li2020">{{Cite journal |last=Li |first=Fay-Wei |display-authors=etal |year=2020 |title=Anthoceros genomes illuminate the origin of land plants and the unique biology of hornworts |journal=Nature Plants |volume=6 |issue=3 |pages=259–272 |doi=10.1038/s41477-020-0618-2 |pmc=8075897 |pmid=32170292 |hdl-access=free |hdl=10261/234303}}</ref><ref name="Sousa2020a">{{Cite journal |last=Sousa |first=Filipe |display-authors=etal |year=2020 |title=The Chloroplast Land Plant Phylogeny: Analyses Employing Better-Fitting Tree- and Site-Heterogeneous Composition Models |journal=Frontiers in Plant Science |volume=11 |page=1062 |doi=10.3389/fpls.2020.01062 |pmc=7373204 |pmid=32760416 |doi-access=free}}</ref><ref name="Su2021">{{Cite journal |last=Su |first=Danyan |display-authors=etal |year=2021 |title=Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants |journal=Molecular Biology and Evolution |volume=38 |issue=8 |pages=3332–3344 |doi=10.1093/molbev/msab106 |pmc=8321542 |pmid=33871608}}</ref> Nevertheless, phylogenies based on mitochondrial sequences fail to support the monophyletic view.<ref name="Sousa2020b">{{Cite journal |last=Sousa |first=Filipe |display-authors=etal |year=2020 |title=The mitochondrial phylogeny of land plants shows support for Setaphyta under composition-heterogeneous substitution models |journal=PeerJ |volume=8 |issue=4 |page=e8995 |doi=10.7717/peerj.8995 |pmc=7194085 |pmid=32377448 |doi-access=free }}</ref>
 The three bryophyte [[clade]]s are the [[Marchantiophyta]] (liverworts), [[moss|Bryophyta]] (mosses) and [[Anthocerotophyta]] (hornworts).<ref name="glossary">{{Cite web |title=GLOSSARY B |url=http://www.emc.maricopa.edu/faculty/farabee/BIOBK/BioBookglossB.html |url-status=dead |archive-url=https://web.archive.org/web/20090402235253/http://www.emc.maricopa.edu/faculty/farabee/biobk/BioBookglossB.html |archive-date=2009-04-02 |access-date=2009-03-26}}</ref> However, it has been proposed that these clades are de-ranked to the classes Marchantiopsida, Bryopsida, and Anthocerotopsida, respectively.<ref name="Sousa2019" /> There is now strong evidence that the liverworts and mosses belong to a monophyletic clade, called [[Setaphyta]].<ref name="Puttick2018" /><ref name=Sousa2020b/><ref name="Cox2018">{{Cite journal |last=Cox |first=Cymon J. |year=2018 |title=Land Plant Molecular Phylogenetics: A Review with Comments on Evaluating Incongruence Among Phylogenies |journal=Critical Reviews in Plant Sciences |volume=37 |issue=2–3 |pages=113–127 |doi=10.1080/07352689.2018.1482443 |hdl-access=free |hdl=10400.1/14557 |s2cid=92198979}}</ref>
 ===Monophyletic view===
-The favoured model, based on amino acids phylogenies, indicates bryophytes as a monophyletic group:<ref name=CoxLiFostEmbl14/>
+The favored model, based on amino acids phylogenies, indicates bryophytes as a monophyletic group:<ref name=CoxLiFostEmbl14/>
 {{Clade|style=line-height:100%;
 |label1=[[embryophyte]]s
 |1={{Clade
+   |1=[[vascular plant]]s
-   |label1=bryophytes
-   |1={{clade
+   |label2=bryophytes
+   |2={{clade
-      |label1=[[Setaphyta|setaphyte]]s
-      |1={{clade
+      |1=[[hornwort]]s
-         |1=[[liverwort]]s
+      |label2=[[Setaphyta|setaphyte]]s
-         |2=[[moss]]es}}
+      |2={{clade
-     |2=[[hornwort]]s
+         |1=[[moss]]es
+         |2=[[liverwort]]s
          }}
+      }}
-   |2=[[vascular plant]]s
    }}
 }}
@@ Line 68: / Line 73: @@
 {{Barlabel
 |size=4
-|at1=2|bar1=green|label1=bryophytes
+|at1=3|bar1=green|label1=bryophytes
 |cladogram=
 {{Clade
 |label1=embryophytes
-|1={{Cladex
+|1={{Clade
+   |1={{Clade
-   |label1=[[Setaphyta|setaphytes]]
-   |1={{Cladex
+      |1=[[vascular plants]]
-       |1=[[liverworts]]|barbegin1=green
+      |2=[[hornworts]]|barbegin2=green
-       |2=[[mosses]]|bar2=green   }}
-   |2={{Cladex
-       |1=[[hornworts]]|barend1=green
-       |2=[[vascular plants]]
-         }}
       }}
+   |label2=[[Setaphyta|setaphytes]]
+   |2={{Clade
+      |1=[[mosses]]|bar1=green
+      |2=[[liverworts]]|barend2=green
+      }}
    }}
+}}
 }}
@@ Line 89: / Line 95: @@
 ===Traditional morphology{{anchor|Taxonomy}}===
-Traditionally, when basing classifications on morphological characters, bryophytes have been distinguished by their lack of vascular structure. However, this distinction is problematic, firstly because some of the earliest-diverging (but now extinct) non-bryophytes, such as the [[horneophyte]]s, did not have true vascular tissue, and secondly because many mosses have well-developed water-conducting vessels.<ref name="Bell">{{Cite journal |last=Bell |first=N. E. |last2=Hyvönen |first2=J. |name-list-style=amp |year=2010 |title=Phylogeny of the moss class Polytrichopsida (BRYOPHYTA): Generic-level structure and incongruent gene trees |url=https://zenodo.org/record/1065524 |journal=Molecular Phylogenetics and Evolution |volume=55 |issue=2 |pages=381–398 |doi=10.1016/j.ympev.2010.02.004 |pmid=20152915}}</ref><ref name="Bodribbetal">{{Cite journal |last=Bodribb |first=T.J. |display-authors=etal |year=2020 |title=Advanced vascular function discovered in a widespread moss |journal=Nature Plants |volume=6 |issue=3 |pages=273–279 |doi=10.1038/s41477-020-0602-x |pmid=32170283 |s2cid=212641738}}</ref> A more useful distinction may lie in the structure of their [[sporophyte]]s. In bryophytes, the sporophyte is a simple unbranched structure with a single spore-forming organ ([[sporangium]]), whereas in all other land plants, the [[polysporangiophyte]]s, the sporophyte is branched and carries many sporangia.<ref name="Kenrick1997a">{{Cite book |last=Kenrick |first=Paul |title=The Origin and Early Diversification of Land Plants: A Cladistic Study |last2=Crane |first2=Peter R. |publisher=Smithsonian Institution Press |year=1997a |isbn=978-1-56098-730-7 |location=Washington, D.C. |name-list-style=amp}}</ref><ref name="Kenrick1997b">{{Cite journal |last=Kenrick |first=P. |last2=Crane |first2=P.R. |name-list-style=amp |year=1997b |title=The origin and early evolution of plants on land |journal=Nature |volume=389 |issue=6646 |pages=33–39 |bibcode=1997Natur.389...33K |doi=10.1038/37918 |s2cid=3866183}}</ref> The contrast is shown in the cladogram below:<ref name="Crane2004">{{Cite journal |last=Crane |first=Peter R. |last2=Herendeen |first2=Patrick |last3=Friis |first3=Else Marie |date=October 2004 |title=Fossils and plant phylogeny |journal=American Journal of Botany |volume=91 |issue=10 |pages=1683–1699 |doi=10.3732/ajb.91.10.1683 |pmid=21652317 |doi-access=free}}</ref>
+Traditionally, when basing classifications on morphological characters, bryophytes have been distinguished by their lack of vascular structure. However, this distinction is problematic, firstly because some of the earliest-diverging (but now extinct) non-bryophytes, such as the [[horneophyte]]s, did not have true vascular tissue, and secondly because many mosses have well-developed water-conducting vessels.<ref name="Bell">{{Cite journal |last1=Bell |first1=N. E. |last2=Hyvönen |first2=J. |name-list-style=amp |year=2010 |title=Phylogeny of the moss class Polytrichopsida (BRYOPHYTA): Generic-level structure and incongruent gene trees |url=https://zenodo.org/record/1065524 |journal=Molecular Phylogenetics and Evolution |volume=55 |issue=2 |pages=381–398 |doi=10.1016/j.ympev.2010.02.004 |pmid=20152915}}</ref><ref name="Bodribbetal">{{Cite journal |last=Bodribb |first=T.J. |display-authors=etal |year=2020 |title=Advanced vascular function discovered in a widespread moss |journal=Nature Plants |volume=6 |issue=3 |pages=273–279 |doi=10.1038/s41477-020-0602-x |pmid=32170283 |s2cid=212641738}}</ref> A more useful distinction may lie in the structure of their [[sporophyte]]s. In bryophytes, the sporophyte is a simple unbranched structure with a single spore-forming organ ([[sporangium]]), whereas in all other land plants, the [[polysporangiophyte]]s, the sporophyte is branched and carries many sporangia.<ref name="Kenrick1997a">{{Cite book |last1=Kenrick |first1=Paul |title=The Origin and Early Diversification of Land Plants: A Cladistic Study |last2=Crane |first2=Peter R. |publisher=Smithsonian Institution Press |year=1997a |isbn=978-1-56098-730-7 |location=Washington, D.C. |name-list-style=amp}}</ref><ref name="Kenrick1997b">{{Cite journal |last1=Kenrick |first1=P. |last2=Crane |first2=P.R. |name-list-style=amp |year=1997b |title=The origin and early evolution of plants on land |journal=Nature |volume=389 |issue=6646 |pages=33–39 |bibcode=1997Natur.389...33K |doi=10.1038/37918 |s2cid=3866183}}</ref> The contrast is shown in the cladogram below:<ref name="Crane2004">{{Cite journal |last1=Crane |first1=Peter R. |last2=Herendeen |first2=Patrick |last3=Friis |first3=Else Marie |date=October 2004 |title=Fossils and plant phylogeny |journal=American Journal of Botany |volume=91 |issue=10 |pages=1683–1699 |doi=10.3732/ajb.91.10.1683 |pmid=21652317 |doi-access=free}}</ref>
 {{clade|style=line-height:100%;
@@ Line 98: / Line 104: @@
 |label1=polysporangiophytes
 |1={{clade
-  |1="protracheophytes", such as ''[[Horneophyton]]'' or ''[[Aglaophyton]]''
+  |1="protracheophytes", such as ''[[Aglaophyton]]'' or ''[[Horneophyton]]''
   |2=tracheophytes or vascular plants
 }}
@@ Line 106: / Line 112: @@
 ==Evolution==
-There have probably been several different terrestrialization events, in which originally aquatic organisms colonized the land, just within the lineage of the [[Viridiplantae]].<ref name="Gerrienne">{{Cite journal |last=Gerrienne |first=Philippe |last2=Servais |first2=Thomas |last3=Vecoli |first3=Marco |date=April 2016 |title=Plant evolution and terrestrialization during Palaeozoic times—The phylogenetic context |journal=Review of Palaeobotany and Palynology |volume=227 |pages=4–18 |doi=10.1016/j.revpalbo.2016.01.004}}</ref> Between 510 - 630 million years ago, however, land plants emerged within the [[green algae]].<ref>{{Cite journal |last=Delwiche |first=Charles F. |last2=Timme |first2=Ruth E. |date=2011-06-07 |title=Plants |url=https://doi.org/10.1016/j.cub.2011.04.021 |journal=Current Biology |volume=21 |issue=11 |pages=R417–R422 |doi=10.1016/j.cub.2011.04.021 |issn=0960-9822 |pmid=21640897 |s2cid=235312105|doi-access=free }}</ref> Molecular phylogenetic studies conclude that bryophytes are the earliest diverging lineages of the extant land plants.<ref name="Konrat2">{{Cite journal |last=Konrat |first=M. |last2=Shaw |first2=A.J. |last3=Renzaglia |first3=K.S. |year=2010 |title=A special issue of ''Phytotaxa'' dedicated to Bryophytes: The closest living relatives of early land plants |journal=Phytotaxa |volume=9 |pages=5–10 |doi=10.11646/phytotaxa.9.1.3 |doi-access=free}}</ref><ref name=pmid12415314/><ref name="KaroArumBoorDuff102">{{Cite journal |last=Karol |first=Kenneth G. |last2=Arumuganathan |first2=Kathiravetpillai |last3=Boore |first3=Jeffrey L. |last4=Duffy |first4=Aaron M. |last5=Everett |first5=Karin DE |last6=Hall |first6=John D. |last7=Hansen |first7=S.K. |last8=Kuehl |first8=Jennifer V. |last9=Mandoli |first9=Dina F. |last10=Mishler |first10=Brent D. |last11=Olmstead |first11=Richard G. |name-list-style=amp |year=2010 |title=Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages |journal=BMC Evolutionary Biology |volume=10 |issue=1 |page=321 |doi=10.1186/1471-2148-10-321 |pmc=3087542 |pmid=20969798 |last12=Renzaglia |first12=Karen S. |last13=Wolf |first13=Paul G.}}</ref><ref name="Shaw">{{Cite journal |last=Shaw |first=A. Jonathan |last2=Szövényi |first2=Péter |last3=Shaw |first3=Blanka |date=March 2011 |title=Bryophyte diversity and evolution: Windows into the early evolution of land plants |journal=American Journal of Botany |volume=98 |issue=3 |pages=352–369 |doi=10.3732/ajb.1000316 |pmid=21613131}}</ref> They provide insights into the migration of plants from aquatic environments to land. A number of physical features link bryophytes to both land plants and aquatic plants.<ref>{{Cite journal |last=Donoghue |first=Philip C. J. |last2=Harrison |first2=C. Jill |last3=Paps |first3=Jordi |last4=Schneider |first4=Harald |date=11 October 2021 |title=The evolutionary emergence of land plants |url=https://research-information.bris.ac.uk/en/publications/the-evolutionary-emergence-of-land-plants |journal=Current Biology |volume=31 |issue=19 |pages=R1281–R1298 |doi=10.1016/j.cub.2021.07.038 |pmid=34637740 |hdl=1983/662d176e-fcf4-40bf-aa8c-5694a86bd41d |s2cid=238588736|doi-access=free }}</ref>
+There have probably been several different terrestrialization events, in which originally aquatic organisms colonized the land, just within the lineage of the [[Viridiplantae]].<ref name="Gerrienne">{{Cite journal |last1=Gerrienne |first1=Philippe |last2=Servais |first2=Thomas |last3=Vecoli |first3=Marco |date=April 2016 |title=Plant evolution and terrestrialization during Palaeozoic times—The phylogenetic context |journal=Review of Palaeobotany and Palynology |volume=227 |pages=4–18 |doi=10.1016/j.revpalbo.2016.01.004}}</ref> Between 510–630 million years ago, however, land plants emerged within the [[green algae]].<ref>{{Cite journal |last1=Delwiche |first1=Charles F. |last2=Timme |first2=Ruth E. |date=2011-06-07 |title=Plants |journal=Current Biology |volume=21 |issue=11 |pages=R417–R422 |doi=10.1016/j.cub.2011.04.021 |issn=0960-9822 |pmid=21640897 |s2cid=235312105|doi-access=free }}</ref> Molecular phylogenetic studies conclude that bryophytes are the earliest diverging lineages of the extant land plants.<ref name="Konrat2"/><ref name=pmid12415314/><ref name="KaroArumBoorDuff102">{{Cite journal |last1=Karol |first1=Kenneth G. |last2=Arumuganathan |first2=Kathiravetpillai |last3=Boore |first3=Jeffrey L. |last4=Duffy |first4=Aaron M. |last5=Everett |first5=Karin DE |last6=Hall |first6=John D. |last7=Hansen |first7=S.K. |last8=Kuehl |first8=Jennifer V. |last9=Mandoli |first9=Dina F. |last10=Mishler |first10=Brent D. |last11=Olmstead |first11=Richard G. |name-list-style=amp |year=2010 |title=Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages |journal=BMC Evolutionary Biology |volume=10 |issue=1 |page=321 |doi=10.1186/1471-2148-10-321 |pmc=3087542 |pmid=20969798 |last12=Renzaglia |first12=Karen S. |last13=Wolf |first13=Paul G. |doi-access=free }}</ref><ref name="Shaw">{{Cite journal |last1=Shaw |first1=A. Jonathan |last2=Szövényi |first2=Péter |last3=Shaw |first3=Blanka |date=March 2011 |title=Bryophyte diversity and evolution: Windows into the early evolution of land plants |journal=American Journal of Botany |volume=98 |issue=3 |pages=352–369 |doi=10.3732/ajb.1000316 |pmid=21613131}}</ref> They provide insights into the migration of plants from aquatic environments to land. A number of physical features link bryophytes to both land plants and aquatic plants.<ref>{{Cite journal |last1=Donoghue |first1=Philip C. J. |last2=Harrison |first2=C. Jill |last3=Paps |first3=Jordi |last4=Schneider |first4=Harald |date=11 October 2021 |title=The evolutionary emergence of land plants |url=https://research-information.bris.ac.uk/en/publications/the-evolutionary-emergence-of-land-plants |journal=Current Biology |volume=31 |issue=19 |pages=R1281–R1298 |doi=10.1016/j.cub.2021.07.038 |pmid=34637740 |hdl=1983/662d176e-fcf4-40bf-aa8c-5694a86bd41d |s2cid=238588736|doi-access=free |hdl-access=free }}</ref>
 === Similarities to algae and vascular plants ===
-Green algae, bryophytes and vascular plants all have chlorophyll a and b, and the chloroplast structures are similar.<ref name="Evert" /> Like green algae and land plants, bryophytes also produce starch stored in the plastids and contain cellulose in their walls.<ref name="Evert" /> Distinct adaptations observed in bryophytes have allowed plants to colonize Earth's terrestrial environments. To prevent desiccation of plant tissues in a terrestrial environment, a waxy cuticle covering the soft tissue of the plant may be present, providing protection. In hornworts and mosses, stomata provide gas exchange between the atmosphere and an internal intercellular space system. The development of gametangia provided further protection specifically for gametes, the zygote and the developing sporophyte.<ref name="Bryophytes">{{Cite web |last=Purcell |first=Adam |title=Bryophytes |url=http://basicbiology.net/plants/lycophytes/bryophytes.php |website=Basic Biology}}</ref> The bryophytes and vascular plants ([[embryophyte]]s) also have embryonic development which is not seen in green algae.<ref name="Evert" /> While bryophytes have no truly vascularized tissue, they do have organs that are specialized for transport of water and other specific functions, analogous for example to the functions of leaves and stems in vascular land plants.<ref name="Evert">{{Cite book |last=Evert |first=Ray |title=Biology of Plants |last2=Eichhorn |first2=Susan |publisher=W.H.Freeman and Company Publishers |year=2013}}</ref>
+Green algae, bryophytes and vascular plants all have chlorophyll a and b, and the chloroplast structures are similar.<ref name="Evert" /> Like green algae and land plants, bryophytes also produce starch stored in the plastids and contain cellulose in their walls.<ref name="Evert" /> Distinct adaptations observed in bryophytes have allowed plants to colonize Earth's terrestrial environments. To prevent desiccation of plant tissues in a terrestrial environment, a waxy cuticle covering the soft tissue of the plant may be present, providing protection. In hornworts and mosses, stomata provide gas exchange between the atmosphere and an internal intercellular space system. The development of gametangia provided further protection specifically for gametes, the zygote and the developing sporophyte.<ref name="Bryophytes">{{Cite web |last=Purcell |first=Adam |title=Bryophytes |url=http://basicbiology.net/plants/lycophytes/bryophytes.php |website=Basic Biology}}</ref> The bryophytes and vascular plants ([[embryophyte]]s) also have embryonic development which is not seen in green algae.<ref name="Evert" /> While bryophytes have no truly vascularized tissue, they do have organs that are specialized for transport of water and other specific functions, analogous for example to the functions of leaves and stems in vascular land plants.<ref name="Evert">{{Cite book |last1=Evert |first1=Ray |title=Biology of Plants |last2=Eichhorn |first2=Susan |publisher=W.H.Freeman and Company Publishers |year=2013}}</ref>
 Bryophytes depend on water for reproduction and survival. In common with ferns and [[Lycopodiopsida|lycophytes]], a thin layer of water is required on the surface of the plant to enable the movement of the flagellated sperm between gametophytes and the fertilization of an egg.<ref name="Bryophytes" />
@@ Line 118: / Line 124: @@
 {| class="wikitable"
 |-
 !
 ! Liverworts
 ! Mosses
@@ Line 158: / Line 164: @@
 {| class="wikitable"
 |-
 !
 ! Liverworts
 ! Mosses
@@ Line 213: / Line 219: @@
 | Present
 |-
 !scope=row| [[Dehiscence (botany)|Dehiscence]]
 | Longitudinal or irregular
 | Transverse
 | Longitudinal
 |}
@@ Line 225: / Line 231: @@
 * Soil Conditioning
 * Bioindicators
 * Moss gardens
 * Pesticides
 Characteristics of bryophytes make them useful to the environment. Depending on the specific plant texture, bryophytes have been shown to help improve the water retention and air space within soil.<ref name="Glime">{{Cite web |last=Glime |first=Janice |title=Economic and Ethnic Uses of Bryophytes |url=http://flora.huh.harvard.edu/FloraData/001/WebFiles/fna27/FNA27-Chapter2.pdf |website=harvard.edu}}</ref> Bryophytes are used in pollution studies to indicate soil pollution (such as the presence of heavy metals), air pollution, and UV-B radiation.<ref name="Glime" /> Gardens in Japan are designed with moss to create peaceful sanctuaries.<ref name="Glime" /> Some bryophytes have been found to produce natural pesticides. The liverwort, ''Plagiochila,'' produces a chemical that is poisonous to mice.<ref name="Glime" /> Other bryophytes produce chemicals that are antifeedants which protect them from being eaten by slugs.<ref name="Glime" /> When ''Phythium sphagnum'' is sprinkled on the soil of germinating seeds, it inhibits growth of "damping off fungus" which would otherwise kill young seedlings.<ref name="Wolffhechel">{{Cite journal |last=Wolffhechel |first=H. |date=April 1988 |title=The suppressiveness of sphagnum peat to ''Pythium'' spp |journal=Acta Horticulturae |issue=221 |pages=217–222 |doi=10.17660/actahortic.1988.221.22}}</ref>
 [[File:Schultz Sphagnum Peat Moss.jpg|thumb|Moss peat is made from ''Sphagnum'']]
@@ Line 244: / Line 250: @@
 ==Bibliography==
-* {{Cite journal |last=Lesica |first=P. |last2=McCune |first2=B. |last3=Cooper |first3=S. V. |last4=Hong |first4=W. S. |year=1991 |title=Differences in lichen and bryophyte communities between old-growth and managed second-growth forests in the Swan Valley, Montana |journal=Canadian Journal of Botany |volume=69 |issue=8 |pages=1745–1755 |doi=10.1139/b91-222}}
+* {{Cite journal |last1=Lesica |first1=P. |last2=McCune |first2=B. |last3=Cooper |first3=S. V. |last4=Hong |first4=W. S. |year=1991 |title=Differences in lichen and bryophyte communities between old-growth and managed second-growth forests in the Swan Valley, Montana |journal=Canadian Journal of Botany |volume=69 |issue=8 |pages=1745–1755 |doi=10.1139/b91-222}}
 == External links ==
 {{commons category|Bryophytes}}
 {{wiktionary}}
-{{Wikisource1911Enc|Bryophyta}}
+{{EB1911 poster|Bryophyta}}
 * [http://www.andrewspink.nl/mosses/ Andrew's Moss Site] Photos of bryophytes
-*27-May-2013 [https://www.bbc.co.uk/news/science-environment-22656239 ''Centuries-old frozen plants revived''], 400-year-old bryophyte specimens left behind by [[Retreat of glaciers since 1850|retreating glaciers]] in Canada are brought back to life in the laboratory.
+* 27-May-2013 [https://www.bbc.co.uk/news/science-environment-22656239 ''Centuries-old frozen plants revived''], 400-year-old bryophyte specimens left behind by [[Retreat of glaciers since 1850|retreating glaciers]] in Canada are brought back to life in the laboratory.
-*{{Cite journal |last=Farge |first=Catherine La |last2=Williams |first2=Krista H. |last3=England |first3=John H. |date=22 May 2013 |title=Regeneration of Little Ice Age bryophytes emerging from a polar glacier with implications of totipotency in extreme environments |journal=Proceedings of the National Academy of Sciences |volume=110 |issue=24 |pages=9839–9844 |bibcode=2013PNAS..110.9839L |doi=10.1073/pnas.1304199110 |pmc=3683725 |pmid=23716658 |doi-access=free}}
+* {{Cite journal |last1=Farge |first1=Catherine La |last2=Williams |first2=Krista H. |last3=England |first3=John H. |date=22 May 2013 |title=Regeneration of Little Ice Age bryophytes emerging from a polar glacier with implications of totipotency in extreme environments |journal=Proceedings of the National Academy of Sciences |volume=110 |issue=24 |pages=9839–9844 |bibcode=2013PNAS..110.9839L |doi=10.1073/pnas.1304199110 |pmc=3683725 |pmid=23716658 |doi-access=free}}
 * Magill, R. E., ed. (1990). ''Glossarium polyglottum bryologiae. A multilingual glossary for bryology''. Monographs in Systematic Botany from the Missouri Botanical Garden, v. 33, 297 pp. [https://web.archive.org/web/20041209031512/http://www.mobot.org/MOBOT/tropicos/most/Glossary/glosefr.html Online version: Internet Archive].

Latest revision as of 08:15, 29 October 2024

Features

Habitat

Life cycle

Sexuality

Classification and phylogeny

Monophyletic view

Paraphyletic view

Traditional morphology

Evolution

Similarities to algae and vascular plants

Comparative morphology

Uses

See also

References

Bibliography

External links