Bryopsida

The Bryopsida constitute the largest class of mosses, containing 95% of all moss species. It consists of approximately 11,500 species, common throughout the whole world.

Bryopsida
Arthrodontous capsule of Dicranella varia
Scientific classification
Kingdom: Plantae
Division: Bryophyta
Subdivision: Bryophytina
Class: Bryopsida
(Limpr.) Rothm.
Subclasses [1]

The group is distinguished by having spore capsules with teeth that are arthrodontous; the teeth are separate from each other and jointed at the base where they attach to the opening of the capsule.[2] These teeth are exposed when the covering operculum falls off. In other groups of mosses, the capsule is either nematodontous with an attached operculum, or else splits open without operculum or teeth.

Morphological groups

The Bryopsida can be simplified into three groups: the acrocarpous (pinnate), the pleurocarpous (side-fruited), and the cladocarpous (branching) mosses. This is based on the position of the perichaetia and sporophytes.

Acrocarps are generally characterized by an upright growth habit that is unbranched or only sparingly branched. Branching is usually sympodial with the branches similar to main shoot where they originate. Branches below the perichaetium are called subfloral innovations.

Pleurocarps are generally characterized by creeping shoot systems and extensive lateral branching. The main stem is indeterminant and offshooting branches may be dissimilar. The perichaetia in pleurocarps are produced at the tips of extremely reduced, basally swollen lateral branches that are morphologically distinct from the vegetative branches.

Cladocarps are mosses which produce perichaetia at the tips of unspecialized lateral branches. Such branches are themselves capable of branching.

Although acrocarps, pleurocarps, and cladocarps generally have different branching habits, it is the morphology of the perichaetia which defines the groups.[3]

Capsule structure

Among the Bryopsida, the structure of the capsule (sporangium) and its pattern of development is very useful both for classifying and for identifying moss families. Most Bryopsida produce a capsule with a lid (the operculum) which falls off when the spores inside are mature and thus ready to be dispersed. The opening thus revealed is called the stoma (meaning "mouth") and is surrounded by one or two peristomes. A peristome is a ring of triangular "teeth" formed from the remnants of specially thickened cell walls. There are usually 16 such teeth in a single peristome, and in the Bryopsida the teeth are separate from each other and able to both fold in to cover the stoma as well as fold back to open the stoma. This articulation of the teeth is termed arthrodontous.

There are two basic arthrodontous peristome types.[4] The first type is termed haplolepidous and consists of a single circle of 16 peristome teeth. This type of peristome is characteristic of subclass Dicranidae. The second type is the diplolepidous peristome found in subclasses Bryidae, Funariidae, and Timmiidae. In this type, there are two rings of peristome teeth—an inner endostome (short for endoperistome) and an exostome. The endostome is a more delicate membrane, and its teeth are aligned between the teeth of the exostome. There are a few mosses in the Bryopsida that have no peristome in their capsules. These mosses still undergo the same cell division patterns in capsule development, but the teeth do not fully develop.

Classification

In the past, the group Bryopsida included all mosses. Current circumscriptions of the group are more limited.[2][5]

class Bryopsida
subclass Buxbaumiidae (only Buxbaumia)
subclass Diphysciidae (only Diphyscium)
subclass Timmiidae (only Timmia)
subclass Funariidae (5 families)
subclass Dicranidae (24 families)
subclass Bryidae (71 families)
 
 

Oedipodiopsida

 
 
 
 

Tetraphidopsida

 
 

Polytrichopsida

 
Bryopsida
 

Buxbaumiidae

 
 
 

Diphysciidae

 
 
 

Timmiidae

 
 

Funariidae

 
 

Dicranidae

 
Bryidae
 

Bryanae (paraphyletic)

 
 

Hypnanae

 
 
 
 
 
 
 
 
The current composition and phylogeny of the Bryopsida.[5][6]

Phylogeny

A detailed phylogeny to the level of order, based on the work by Novíkov & Barabaš-Krasni 2015.[7]

 
Buxbaumiidae

Buxbaumiales

 
 
Diphysciidae

Diphysciales

 
 
Funariidae
 

Gigaspermales

 
 

Encalyptales

 
 

Funariales

 
 
 
Timmiidae

Timmiales

 
 
Dicranidae
 
 

Archidiales

 
 

Scouleriales

 
 

Grimmiales

 
 
 
 

Bryoxiphiales

 
 

Pottiales

 
 

Dicranales

 
 
 
Bryidae
 
 

Bartramiales

 
 

Hedwigiales

 
 
 

Splachnales

 
 
 

Bryales

 
 
 

Orthotrichales

 
 
 

Orthodontiales

 
 
 

Rhizogoniales

 
 
 

Aulacomniales

 
Hypnanae
 

Hypnodendrales

 
 
 

Ptychomniales

 
 
 

Hookeriales

 
 

Hypnales

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

References

  1. Buck, William R. & Bernard Goffinet. (2000) "Morphology and classification of mosses", pages 71-123 in A. Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology. (Cambridge: Cambridge University Press). ISBN 0-521-66097-1
  2. http://flora.huh.harvard.edu/FloraData/001/WebFiles/fna27/FNA27-Chapter1.pdf
  3. Edwards, S. R. 1984. "Homologies and inter-relationships of moss peristomes", pages 658-695 in R. M. Schuster (Ed.) New Manual of Bryology. (Japan: The Hattori Botanical Laboratory). ISBN 49381633045.
  4. Goffinet, B.; Buck, W. R.; Shaw, A. J. (2008). "Morphology and Classification of the Bryophyta". In Goffinet, B.; Shaw, J. (eds.). Bryophyte Biology (2nd ed.). New York: Cambridge University Press. pp. 55–138. ISBN 978-0-521-87225-6.
  5. Goffinet, Bernard; William R. Buck (2004). "Systematics of the Bryophyta (Mosses): From molecules to a revised classification". Monographs in Systematic Botany. Molecular Systematics of Bryophytes. Missouri Botanical Garden Press. 98: 205–239. ISBN 1-930723-38-5.
  6. Novíkov & Barabaš-Krasni (2015). Modern plant systematics. Liga-Pres. p. 685. doi:10.13140/RG.2.1.4745.6164. ISBN 978-966-397-276-3.
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