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Hornwort

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This article is about the non-vascular plants. The name Hornwort also refers to aquatic plants of the genus Ceratophyllum, in the family Ceratophyllaceae.

Hornwort
Temporal range: 90–0 Ma Upper Cretaceous (but see text) to recent
Phaeoceros laevis (L.) Prosk.
Scientific classification
Kingdom:
Plantae
Division:
Anthocerotophyta

Stotler & Stotl.-Crand.[1]
Classes & Orders
Leiosporocerotopsida
Anthocerotopsida

see Classification.

Synonyms

Anthocerotae

Hornworts are a group of bryophytes, or non-vascular plants, comprising the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. The flattened, green plant body of a hornwort is the gametophyte plant.

Hornworts may be found worldwide, though they tend to grow only in places that 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.

Description

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 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 that both manufactures and stores food. This particular feature is very unusual in land plants, but is common among algae.

Many hornworts develop internal mucilage-filled cavities when groups of cells break down. These cavities are invaded by photosynthetic cyanobacteria, 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 pores on the underside of the thallus. These pores superficially resemble the stomata of other plants.

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 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.

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 and pseudo-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 the proximal surface, and with a distal surface ornamented with bumps or spines.

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.[2] 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 with stomata.[3]

Classification

The hornwort Dendroceros crispus growing on the bark of a tree.

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. The division Bryophyta is now restricted to include only mosses.

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 containing a single family.

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.

Phylogeny

Recent studies of molecular, ultrastructural, and morphological data have yielded a new classification of hornworts.[4]

order Leiosporocerotales
Leiosporocerotaceae

order Anthocerotales

Anthocerotaceae

order Notothyladales

Notothyladaceae

order Phymatocerotales

Phymatocerotaceae

order Dendrocerotales

Dendrocerotaceae
The current phylogeny and composition of the Anthocerotophyta.[4][5]

See also

References

  1. ^ Stotler, Raymond E. (1977). "A checklist of the liverworts and hornworts of North America". The Bryologist. 80 (3). American Bryological and Lichenological Society: 405–428. doi:10.2307/3242017. JSTOR 3242017. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  2. ^ Qiu, Y.L. (2006). "The deepest divergences in land plants inferred from phylogenomic evidence". Proceedings of the National Academy of Sciences. 103 (42): 15511–6. doi:10.1073/pnas.0603335103. PMC 1622854. PMID 17030812. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: extra punctuation (link)
  3. ^ Kenrick, Paul (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. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  4. ^ a b Duff, R. Joel (2007). "Progress 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. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  5. ^ Villareal, J. C. "A synthesis of hornwort diversity: Patterns, causes and future work" (pdf). Phytotaxa. 9: 150–166. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  • 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. Vol. 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.
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