Hornwort

Hornworts are a group of non-vascular plants constituting the division

Anthocerotophyta. The common name refers to the elongated horn-like
Hornworts
structure, which is the sporophyte. As in mosses and liverworts, the flattened, Temporal range: 90–0 Ma
green plant body of a hornwort is thegametophyte plant. PreЄ Є O S D C P T J K PgN
Upper Cretaceous (but see text) to
Hornworts may be found worldwide, though they tend to grow only in places that
present
are damp or humid. Some species grow in large numbers as tiny weeds in the soil
of gardens and cultivated fields. Large tropical and sub-tropical species of
Dendroceros may be found growing on the bark of trees.

The total number of species is still uncertain. While there are more than 300
published species names, the actual number could be as low as 100-150
species.[2]

Contents
Description Phaeoceros laevis (L.) Prosk.
Life cycle Scientific classification
Evolutionary history
Kingdom: Plantae
Classification
Phylogeny Clade: Embryophytes
See also Division: Anthocerotophyta
References Stotler & Stotl.-Crand.,
External links 1977 [1]

Classes & Orders

Description
Leiosporocerotopsida
The plant body of a hornwort is a haploid gametophyte stage. This stage usually
grows as a thin rosette or ribbon-like thallus between one and five centimeters in Leiosporocerotales
diameter. Each cell of the thallus usually contains just one chloroplast. In most
species, this chloroplast is fused with other organelles to form a large pyrenoid Anthocerotopsida
that both manufactures and stores food. This particular feature is very unusual in
land plants, but is common amongalgae. Anthocerotales
Many hornworts develop internal mucilage-filled cavities when groups of cells Dendrocerotales
break down. These cavities are invaded by photosynthetic cyanobacteria, Notothyladales
especially species of Nostoc. Such colonies of bacteria growing inside the thallus
give the hornwort a distinctive blue-green color. There may also be small slime
Phymatocerotales
pores on the underside of the thallus. These pores superficially resemble the see Classification.
stomata of other plants.
Synonyms
The horn-shaped sporophyte grows from an archegonium embedded deep in the
gametophyte. The sporophyte of a hornwort is unusual in that it grows from a Anthocerotae
meristem near its base, instead of from its tip the way other plants do. Unlike
liverworts, most hornworts have true stomata on their sporophyte as mosses do. The exceptions are the genera Notothylas and
Megaceros, which do not have stomata. The sporophyte of most hornworts are also photosynthetic, which is not the case with
liverworts.[3]

When the sporophyte is mature, it has a multicellular outer layer, a central rod-like columella running up the center, and a layer of
tissue in between that produces spores andpseudo-elaters. The pseudo-elaters are multi-cellular, unlike the elaters of liverworts. They
have helical thickenings that change shape in response to drying out; they twist and thereby help to disperse the spores. Hornwort
spores are relatively large for bryophytes, measuring between 30 and 80 µm in diameter or more. The spores are polar, usually with a
distinctive Y-shaped tri-radiate ridge on theproximal surface, and with a distal surface ornamented with bumps or spines

Life cycle
The life of a hornwort starts from a haploid spore. In most species, there is a single cell inside the spore, and a slender extension of
this cell called the germ tube germinates from the proximal side of the spore.[4] The tip of the germ tube divides to form an octant
(solid geometry) of cells, and the first rhizoid grows as an extension of the original germ cell. The tip continues to divide new cells,
which produces a thalloid protonema. By contrast, species of the family Dendrocerotaceae may begin dividing within the spore,
becoming multicellular and even photosynthetic before the spore germinates.[4] In either case, the protonema is a transitory stage in
the life of a hornwort.

From the protonema grows the adult gametophyte, which is the

persistent and independent stage in the life cycle. This stage usually
grows as a thin rosette or ribbon-like thallus between one and five
centimeters in diameter, and several layers of cells in thickness. It is
green or yellow-green from the chlorophyll in its cells, or bluish-
green when colonies of cyanobacteria grow inside the plant.

When the gametophyte has grown to its adult size, it produces the
sex organs of the hornwort. Most plants are monoecious, with both
sex organs on the same plant, but some plants (even within the same
species) are dioecious, with separate male and female gametophytes.
The female organs are known as archegonia (singular archegonium)
and the male organs are known as antheridia (singular antheridium).
Both kinds of organs develop just below the surface of the plant and
are only later exposed by disintegration of the overlying cells.

The biflagellate sperm must swim from the antheridia, or else be

Life cycle of a typical hornwortPhaeoceros. Click
on the image to enlarge.
three basic regions of the sporophyte.
splashed to the archegonia. When this happens, the sperm and egg
cell fuse to form a zygote, the cell from which the sporophyte stage
of the life cycle will develop. Unlike all other bryophytes, the first
cell division of the zygote is longitudinal. Further divisions produce

At the bottom of the sporophyte (closest to the interior of the gametophyte), is a foot. This is a globular group of cells that receives
nutrients from the parent gametophyte, on which the sporophyte will spend its entire existence. In the middle of the sporophyte (just
above the foot), is a meristem that will continue to divide and produce new cells for the third region. This third region is the capsule.
Both the central and surface cells of the capsule are sterile, but between them is a layer of cells that will divide to produce pseudo-
elaters and spores. These are released from the capsule when it splits lengthwise from the tip.

Evolutionary history
While the fossil record of crown group hornworts only begins in the upper Cretaceous, the lower Devonian Horneophyton may
represent a stem group to the clade, as it possesses a sporangium with central columella not attached at the roof.[5] However, the
same form of columella is also characteristic of basal moss groups, such as the Sphagnopsida and Andreaeopsida, and has been
interpreted as a character common to all early land plants withstomata.[6]

Classification
Hornworts were traditionally considered a class within the division Bryophyta
(bryophytes). However, it now appears that this former division is paraphyletic, so the
hornworts are now given their own division, Anthocerotophyta (sometimes misspelled
Anthocerophyta). The division Bryophyta is now restricted to include onlymosses.

Traditionally, there is a single class of hornworts, called Anthocerotopsida, or older

Anthocerotae. More recently, a second class Leiosporocertotopsida has been segregated
for the singularly unusual species Leiosporoceros dussii. All other hornworts remain in
the class Anthocerotopsida. These two classes are divided further into five orders, each The hornwort Dendroceros
containing a single family. crispus growing on the bark of a
tree.
Among land plants, hornworts appear to be one of the oldest surviving lineages;
cladistic analysis implies that the group originated prior to the Devonian, around the
same time as the mosses and liverworts. There are only about 100 species known, but new species are still being discovered. The
number and names of genera are a current matter of investigation, and several competing classification schemes have been published
since 1988.

Structural features that have been used in the classification of hornworts include: the anatomy of chloroplasts and their numbers
within cells, the presence of a pyrenoid, the numbers of antheridia within androecia, and the arrangement of jacket cells of the
antheridia.[7]

Phylogeny
on of hornworts.[8]
Recent studies of molecular, ultrastructural, and morphological data have yielded a new classificati

order Leiosporocerotales Leiosporocerotaceae Leiosporoceros

Leiosporocerotaceae
Folioceros
Leiosporoceros (1 species)

order Anthocerotales
Anthocerotaceae Sphaerosporoceros

Anthocerotaceae Anthoceros
Anthoceros (ca. 83 species)
Folioceros (17 species) Notothylas
Sphaerosporoceros (2 species)
Notothyladaceae
Phaeoceros
order Notothyladales
Phymatocerotaceae Phymatoceros
Notothyladaceae
Notothylas (21 species) Dendrocerotaceae Phaeomegaceros
Phaeoceros (ca. 41 species)
Paraphymatoceros (1-2 Nothoceros
species)
Hattorioceros (1 species)
Megaceros
Mesoceros (2 species)
order Phymatocerotales Dendroceros

Phymatocerotaceae
Phymatoceros (2 species)

order Dendrocerotales

Dendrocerotaceae
Dendroceros (43 species)
Megaceros (8 species)
Nothoceros (7 species)
Phaeomegaceros (7 species)

[8][9]
The current phylogeny and composition of the Anthocerotophyta.

See also
Embryophyte

References
1. Stotler, Raymond E.; Barbara J. Candall-Stotler (1977). "A checklist of the liverworts and hornworts of North
America". The Bryologist. American Bryological and Lichenological Society . 80 (3): 405–428. doi:10.2307/3242017
(https://doi.org/10.2307%2F3242017). JSTOR 3242017 (https://www.jstor.org/stable/3242017).
2. What is a hornwort? - Australian National Botanic Gardens(https://www.anbg.gov.au/bryophyte/what-is-hornwort.htm
l)
3. Mosses, Liverworts, and Hornworts(http://www.mhhe.com/biosci/genbio/tlw3/eBridge/Chp16/16_2.pdf)
4. Chopra, R. N.; Kumra, P. K. (1988). Biology of Bryophytes. New York: John Wiley & Sons.ISBN 0-470-21359-0.
5. Qiu, Y.L.; Li, L.; Wang, B.; Chen, Z.; Knoop, V.; Groth-malonek, M.; Dombrovska, O.; Lee, J.; Kent, L.; Rest, J.;et al.
(2006). "The deepest divergences in land plants inferred from phylogenomic evidence" (https://www.ncbi.nlm.nih.go
v/pmc/articles/PMC1622854). Proceedings of the National Academy of Sciences . 103 (42): 15511–6.
doi:10.1073/pnas.0603335103(https://doi.org/10.1073%2Fpnas.0603335103) . PMC 1622854 (https://www.ncbi.nlm.
nih.gov/pmc/articles/PMC1622854). PMID 17030812 (https://www.ncbi.nlm.nih.gov/pubmed/17030812).
6. Kenrick, Paul; Peter R. Crane (1997).The Origin and Early Diversification of Land Plants: A Cladistic Study
.
Washington, D. C.: Smithsonian Institution Press. pp. 55–56. ISBN 1-56098-730-8.
7. D. Christine Cargill, Karen S. Renzaglia, Juan Carlos Villarreal, and R. Joel Duff (2005), "Generic concepts within
hornworts: Historical review, contemporary insights and future directions",Australian Systematic Botany, 18: 7–16,
doi:10.1071/sb04012 (https://doi.org/10.1071%2Fsb04012)
8. Duff, R. Joel; Juan Carlos Villarreal; D. Christine Cargill; Karen S. Renzaglia (2007). "Prog
ress and challenges
toward a phylogeny and classification of the hornworts".The Bryologist. 110 (2): 214–243. doi:10.1639/0007-
2745(2007)110[214:PACTDA]2.0.CO;2 (https://doi.org/10.1639%2F0007-2745%282007%29110%5B214%3AP ACT
DA%5D2.0.CO%3B2).
9. Villareal, J. C.; Cargill, D. C.; Hagborg, A.; Söderström, L.; Renzaglia, K. S. (2010)."A synthesis of hornwort
diversity: Patterns, causes and future work"(http://mapress.com/phytotaxa/content/2010/f/pt00009p166.pdf)(pdf).
Phytotaxa. 9: 150–166.

Grolle, Riclef (1983). "Nomina generica Hepaticarum; references, types and synonymies". Acta Botanica Fennica.
121: 1–62.
Hasegawa, J. (1994). "New classification of Anthocerotae".Journal of the Hattori Botanical Laboratory. 76: 21–34.
Renzaglia, Karen S. (1978). "A comparative morphology and developmental anatomy of the Anthocerotophyta".
Journal of the Hattori Botanical Laboratory. 44: 31–90.
Renzaglia, Karen S. & Vaughn, Kevin C. (2000). Anatomy, development, and classification of hornworts. In A.
Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology, pp. 1–20. Cambridge:Cambridge University Press.
ISBN 0-521-66097-1.
 Schofield, W. B. (1985). Introduction to Bryology. New York: Macmillan.
Schuster, Rudolf M. (1992). The Hepaticae and Anthocerotae of North America, East of the Hundredth Meridian . VI.
Chicago: Field Museum of Natural History.
Smith, Gilbert M. (1938).Cryptogamic Botany, Volume II: Bryophytes and Pteridophytes. New York: McGraw-Hill
Book Company.
Watson, E. V. (1971). The Structure and Life of Bryophytes(3rd ed.). London: Hutchinson University Library
. ISBN 0-
09-109301-5.

External links
Hornwort Web Portal
Anthocerotophyta description and fossil history at UCMP
Classification of the Anthocerotophyta
New Zealand Anthocerotae

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