Acrocarpous
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The Bryopsida constitute the largest
class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
of
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
es, containing 95% of all moss
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. It consists of approximately 11,500 species, common throughout the whole world. 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.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). Consequently, mosses in the Class Bryopsida are commonly known as the “joint-toothed” or “arthrodontous” mosses. 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 A sporophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a diploid multicellular organism which produces asexual spores. This stage alternates with a multicellular haploid gametophyte pha ...
. Acrocarps are generally characterized by an upright growth habit that is unbranched or only sparingly branched. Branching is usually
sympodial In botany, sympodial growth is a bifurcating branching pattern where one branch develops more strongly than the other, resulting in the stronger branches forming the primary shoot and the weaker branches appearing laterally. A sympodium, als ...
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.


Capsule structure

Among the Bryopsida, the structure of the capsule (
sporangium A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
) 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
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s 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 Peristome (from the Greek language, Greek ''peri'', meaning 'around' or 'about', and ''stoma'', 'mouth') is an anatomical feature that surrounds an opening to an organ or structure. Some plants, fungi, and shelled gastropods have peristomes. In mo ...
is a ring of triangular "teeth" formed from the remnants of specially thickened
cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...
s. 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. 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 Dicranidae are a widespread and diverse subclass of mosses in class Bryopsida, with many species of dry or disturbed areas. They are distinguished by their spores; the peristome teeth are haplolepideous with a 4:2:3 formula, and an exostome ...
. The second type is the diplolepidous peristome found in subclasses
Bryidae Bryidae is an important subclass of Bryopsida. It is common throughout the whole world. Members have a double peristome with alternating tooth segments. Classification The classification of the Bryidae.Buck, William R. & Bernard Goffinet. 2000. ...
,
Funariidae The Funariidae are a widespread group of mosses in class Bryopsida.Buck, William R. & Bernard Goffinet. 2000. "Morphology and classification of mosses", pages 71-123 ''in'' A. Jonathan Shaw & Bernard Goffinet (Eds.), ''Bryophyte Biology''. (Cambr ...
, and
Timmiidae ''Timmia'' is a genus of moss. It is the only genus in the family Timmiaceae and order Timmiales.Buck, William R. & Bernard Goffinet. 2000. "Morphology and classification of mosses", pages 71-123 ''in'' A. Jonathan Shaw & Bernard Goffinet (Eds.) ...
. 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.


Phylogeny

A detailed phylogeny to the level of order, based on the work by Novíkov & Barabaš-Krasni 2015; Cole, Hilger & Goffinet 2021; Fedosov et al. 2016; Ignatov, Fedosov & Fedorova 2016; Bechteler et al. 2023. Unassigned Dicranidae: * Pseudoditrichales


References

{{Taxonbar, from=Q831482 Plant classes