Overview

Brief Summary

Introduction

More than 95% of moss species belong to the Bryopsida. Diversity in this group has been classified into 90 to 110 families and 11 to 16 orders. The Dicranales and Hypnales are the most diverse groups with 13 and 44 families, respectively.

The most important characteristic of the Bryopsida is the architecture of the ring of teeth (peristome) surrounding the mouth of the sporophyte capsule (Figure 1). Bryopsida are the only mosses that have an arthrodontous peristome, i. e., a peristome in which the teeth are structured by articulated cell wall remnants (see Characteristics).

Figure 1. Closeup of the peristome of the capsule of Sematophyllum (Hypnales). This arthrodontous peristome is composed of a double ring of teeth. The external teeth (exostome) are shorter and thicker than the segments of the internal row (endostome). Note that each tooth in the exostome is alternated with a segment of the endostome.
Image copyright © 2000, Efrain De Luna.

Despite all orders sharing a basic arthrodontous peristome, the Bryopsida comprises a diverse set of species of various gametophytic morphologies. Branching systems combine pinnate (monopodial) and stepwise extension of new branches (sympodial architectures) with apical (acrocarpous) or lateral (cladocarpous and pleurocarpous) sporophyte locations. Leaf cells are variously differentiated forming specialized groups at allar, basal, medial, upper and apical leaf zones. Leaves in some taxa are multilayered stratose, at least in the apex and margins. The sporangia also show a wide range of variation in shape and size of the capsule, operculum, and peristome organization. The outer surface of teeth can be variously ornamented with papillae, striations, or both.

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Comprehensive Description

Characteristics

All of the features of the gametophyte (protonema, gametophore, gametangia) and sporophyte (seta, sporangium and peristome) described for the Bryophyta are applicable to the Bryopsida. Here, the basic patterns of variation in the peristome requires special examination, since it provides most of the characters for the classification at the ordinal level in the Bryopsida.

Each tooth of the arthrodontous peristome type is composed of periclinal (tangential) cell wall remnants between two of the three concentric peristomial cell layers: the outer, primary, and inner peristomial layers (OPL, PPL, and IPL; Figure 2A). Often, a double ring of teeth is present surrounding the capsule mouth (Figure 1). The external ring of teeth is formed by the tangential walls between the OPL and PPL; it is collectively known as the exostome. The internal row consists of the cell wall remnants located between the cell rings of the PPL and IPL, therefore the row of segments is known as endostome.

Depending on the number of cell columns articulated to form the length of a tooth, two primary types of arthrodontous peristomes are known: diplolepidous and haplolepidous.

In diplolepidous peristomes two columns of adjacent OPL cells participate in the formation of an exostome tooth (Figure 2B, D, E). A medial vertical line marking the two cells can be observed along the outer surface of each tooth. This type of peristome can be observed in taxa of the Funariales, Bryales and Hypnales, among others. In haplolepidous peristomes the exostome is missing, but the endostome is well developed and strongly thickened. The outer surface of an endostome segment is made up of just one column of cells from the PPL (Figure 2C). The haplolepidous peristome is characteristic of the Grimmiales, Seligeriales, Archidiales, Pottiales, and Dicranales.

Figure 2. The different peristome types of Bryopsida.
A. Cross section of an embryonic sporophyte capsule.
B. Diplolepidous opposite peristome (Funaria type).
C. Haplolepidous peristome (Dicranum type).
D. Diplolepidous alternate (Orthotrichum type).
E. Diplolepidous alternate (Bryum type).
Drawings copyright © 2000 Efrain de Luna.

Two basic peristome "types" have long been recognized in the diplolepidous mosses: diplolepidous-opposite, and diplolepidous-alternate. In diplolepidous peristomes usually both the exostome and the endostome are present, but in several families the endostome is lacking or reduced. When the two concentric rows are present, each external tooth can be directly opposite or alternated to the internal segments of the endostome.

In the diplolepidous-opposite or "Funaria type", the anticlinal walls of the IPL and PPL are almost perfectly aligned (co-radial), so the endostome segments and the exostome teeth are opposite each other (Figure 2B). This peristome organization in found in the Funariales.

In the diplolepidous-alternate or "Bryum type", anticlinal cell divisions in the IPL are asymmetric, producing narrow cells adjacent to large cells. Later in development, the IPL ring shifts displacing the position of radial IPL cell walls relative to those in the PPL cells. This results in the placement of the endostome segments alternating with the exostome teeth (Figure 2D, E). The diplolepidous alternate peristome is common in the Bryales, Rhizogoniales, Hookeriales, and Hypnales.

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Ecology

Associations

Plant / associate
Acalypta brunnea is associated with Bryopsida

Plant / associate
Acalypta carinata is associated with Bryopsida

Plant / associate
Acalypta nigrina is associated with Bryopsida

Plant / associate
Acalypta parvula is associated with Bryopsida

Plant / associate
Acalypta platycheila is associated with Bryopsida

In Great Britain and/or Ireland:
Plant / associate
fruitbody of Arrhenia acerosa is associated with gametophyte of Bryopsida

Plant / associate
fruitbody of Arrhenia chlorocyanea is associated with gametophyte of Bryopsida

Plant / epiphyte
fruitbody of Arrhenia latispora grows on gametophyte of Bryopsida

Plant / epiphyte
fruitbody of Arrhenia lobata grows on gametophyte of Bryopsida

Plant / associate
fruitbody of Arrhenia obatra is associated with gametophyte of Bryopsida

Foodplant / parasite
fruitbody of Arrhenia retiruga parasitises gametophyte of Bryopsida

Plant / associate
fruitbody of Arrhenia rickenii is associated with gametophyte of Bryopsida

Plant / associate
fruitbody of Arrhenia rustica is associated with gametophyte of Bryopsida

Plant / epiphyte
fruitbody of Arrhenia spathulata grows on gametophyte of Bryopsida

Plant / associate
apothecium of Ascophanus globosopulvinatus is associated with live gametophyte of Bryopsida

Plant / epiphyte
fruitbody of Athelia fibulata grows on Bryopsida

Plant / grows inside
ascoma of Bryomyces microcarpus grows inside leaf of gametophyte of Bryopsida

Plant / associate
basidiome of Camarophyllopsis foetens is associated with Bryopsida
Other: minor host/prey

Plant / associate
Campylosteira verna is associated with Bryopsida

Plant / associate
basidiome of Ceratellopsis sagittiformis is associated with Bryopsida

Plant / associate
basidiome of Cheimonophyllum candidissimum is associated with Bryopsida
Other: minor host/prey

Foodplant / parasite
fruitbody of Chromocyphella muscicola parasitises live gametophyte of Bryopsida

Plant / associate
fruitbody of Clavulinopsis luteonana is associated with Bryopsida

Plant / associate
fruitbody of Clavulinopsis rufipes is associated with Bryopsida

Plant / associate
fruitbody of Clitocybe barbularum is associated with Bryopsida

Plant / associate
fruitbody of Collybia acervata is associated with Bryopsida

Plant / associate
fruitbody of Collybia putilla is associated with Bryopsida

Plant / associate
sclerotium of Collybia tuberosa is associated with Bryopsida

Plant / associate
fruitbody of Conocybe sulcatipes is associated with Bryopsida

Plant / associate
fruitbody of Coprinus cinnamomeotinctus is associated with Bryopsida

Plant / grows among
sporangium of Craterium muscorum grows among gametophyte of Bryopsida

Plant / associate
fruitbody of Cystoderma amianthinum is associated with gametophyte of Bryopsida

Plant / associate
fruitbody of Cystoderma carcharias is associated with gametophyte of Bryopsida
Other: minor host/prey

Plant / associate
fruitbody of Dendrocollybia racemosa is associated with Bryopsida

Plant / associate
apothecium of Discinella menziesii is associated with Bryopsida

Foodplant / sap sucker
nymph of Drymus brunneus sucks sap of Bryopsida

Foodplant / sap sucker
adult of Drymus sylvaticus sucks sap of Bryopsida

Plant / associate
fruitbody of Entoloma anatinum is associated with live Bryopsida

Plant / associate
fruitbody of Entoloma caeruleum is associated with live Bryopsida

Plant / associate
fruitbody of Entoloma favrei is associated with live Bryopsida

Plant / associate
fruitbody of Entoloma inutile is associated with live Bryopsida

Plant / associate
fruitbody of Entoloma lanuginosipes is associated with Bryopsida

Plant / associate
fruitbody of Entoloma olorinum is associated with live Bryopsida

Plant / associate
fruitbody of Entoloma poliopus var. poliopus is associated with Bryopsida

Plant / associate
fruitbody of Entoloma sericellum is associated with Bryopsida

Foodplant / parasite
Eocronartium muscicola parasitises live Bryopsida

Plant / epiphyte
solitary, superficial ascoma of Epibryon bryophilum grows on dying gametophyte of Bryopsida

Plant / epiphyte
superficial ascoma of Epibryon casaresii grows on live gametophyte of Bryopsida

Plant / associate
sporangium of Fuligo muscorum is associated with terrestrial gametophyte of Bryopsida

Plant / associate
fruitbody of Galerina atkinsoniana is associated with Bryopsida

Plant / associate
fruitbody of Galerina calyptrata is associated with Bryopsida

Plant / associate
fruitbody of Galerina cephalotricha is associated with Bryopsida

Plant / associate
fruitbody of Galerina cerina var. cerina is associated with Bryopsida

Plant / associate
fruitbody of Galerina clavata is associated with Bryopsida

Plant / associate
fruitbody of Galerina embolus is associated with Bryopsida

Plant / associate
fruitbody of Galerina hypnorum is associated with Bryopsida

Plant / associate
fruitbody of Galerina jaapii is associated with Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Galerina laevis is associated with Bryopsida

Plant / associate
fruitbody of Galerina mniophila is associated with live Bryopsida

Plant / associate
fruitbody of Galerina praticola is associated with Bryopsida

Plant / associate
fruitbody of Galerina pseudocerina is associated with Bryopsida

Plant / associate
fruitbody of Galerina pseudomniophila is associated with Bryopsida

Plant / associate
fruitbody of Galerina pseudomycenopsis is associated with Bryopsida

Plant / associate
fruitbody of Galerina pumila is associated with Bryopsida

Plant / associate
fruitbody of Galerina subannulata is associated with Bryopsida

Plant / associate
fruitbody of Galerina subcerina is associated with Bryopsida

Plant / associate
fruitbody of Galerina subclavata is associated with Bryopsida

Plant / associate
fruitbody of Galerina uncialis is associated with Bryopsida

Plant / associate
fruitbody of Galerina viscidula is associated with Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Galerina vittiformis is associated with Bryopsida

Plant / associate
fruitbody of Gerronema prescotii is associated with Bryopsida

Plant / associate
fruitbody of Gerronema stevensonii is associated with Bryopsida

Plant / associate
sporocarp of Glomus fuegianum is associated with Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Gymnopilus fulgens is associated with Bryopsida

Foodplant / parasite
solitary apothecium of Helotium fulvum parasitises Bryopsida

Plant / associate
fruitbody of Hohenbuehelia longipes is associated with Bryopsida

Plant / associate
fruitbody of Hygrocybe cantharellus is associated with live Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Hygrocybe citrinopallida is associated with live Bryopsida

Plant / associate
fruitbody of Hygrocybe glutinipes var. glutinipes is associated with live Bryopsida

Plant / associate
fruitbody of Hygrocybe turunda is associated with live Bryopsida
Other: unusual host/prey

Foodplant / parasite
solitary apothecium of Hymenoscyphus phascoides parasitises live leaf of Bryopsida

Plant / associate
gymnothecium of Illosporium curreyi is associated with living Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Inocybe glabrodisca is associated with Bryopsida

Plant / grows among
gregarious apothecium of Lamprospora annulata grows among gametophyte of Bryopsida

Plant / grows among
apothecium of Lamprospora astroidea grows among Bryopsida

Plant / grows among
gregarious apothecium of Lamprospora carbonicola grows among gametophyte of Bryopsida

Plant / grows among
apothecium of Lamprospora crouanii grows among Bryopsida

Plant / grows among
solitary or gregarious apothecium of Lamprospora dictydiola grows among gametophyte of Bryopsida

Foodplant / parasite
fruitbody of Leptosporomyces raunkiaeri parasitises live plant of Bryopsida

Plant / grows among
sessile apothecium of Leucoscypha erminea grows among gametophyte of Bryopsida
Remarks: season: 6
Other: minor host/prey

Plant / grows among
apothecium of Leucoscypha leucotricha grows among gametophyte of Bryopsida
Other: minor host/prey

Plant / resting place / on
Botrydina anamorph of Lichenomphalia umbellifera may be found on plant of Bryopsida
Other: minor host/prey

Plant / associate
puparium of Limnophora maculosa is associated with partially submerged Bryopsida

Plant / grows among
gregarious apothecium of Marcelleina persoonii grows among gametophyte of Bryopsida

Plant / grows among
apothecium of Marcelleina rickii grows among gametophyte of Bryopsida

Plant / grows among
apothecium of Microglossum viride grows among gametophyte of Bryopsida
Other: minor host/prey

Plant / associate
fruitbody of Mycena corynephora is associated with corticolous Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Mycena hiemalis is associated with live colony of Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Mycena latifolia is associated with Bryopsida

Plant / associate
fruitbody of Mycena meliigena is associated with living Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Mycena mirata is associated with living Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Mycena olida is associated with live colony of Bryopsida
Other: major host/prey

Plant / associate
fruitbody of Mycena pseudocorticola is associated with live colony of Bryopsida

Foodplant / saprobe
fruitbody of Mycocalia minutissima is saprobic on wet plant of Bryopsida

Plant / associate
fruitbody of Naucoria amarescens is associated with Bryopsida
Other: minor host/prey

Plant / epiphyte
apothecium of Octospora convexula grows on protonema of Bryopsida

Plant / grows among
apothecium of Octospora melina grows among gametophyte of Bryopsida
Other: minor host/prey

Plant / associate
fruitbody of Omphaliaster asterosporus is associated with Bryopsida

Plant / associate
fruitbody of Omphalina galericolor var. galericolor is associated with Bryopsida

Plant / associate
fruitbody of Omphalina galericolor var. lilacinicolor is associated with Bryopsida

Plant / associate
fruitbody of Omphalina pyxidata is associated with Bryopsida

Plant / associate
fruitbody of Omphalina subhepatica is associated with Bryopsida

Plant / associate
fruitbody of Omphalina wallacei is associated with Bryopsida

Plant / grows among
apothecium of Pezoloma ciliifera grows among gametophyte of Bryopsida

Plant / resting place / under
puparium of Phaonia mystica may be found under Bryopsida

Plant / associate
sporangium of Physarum virescens is associated with terrestrial gametophyte of Bryopsida

Plant / associate
annual, resupinate, drying dark red fruitbody of Physisporinus sanguinolentus is associated with Bryopsida
Other: unusual host/prey

Plant / associate
fruitbody of Pseudoclitocybe obbata is associated with Bryopsida

Plant / associate
apothecium of Pulvinula convexella is associated with gametophyte of Bryopsida
Other: minor host/prey

Plant / epiphyte
thallus of Pyrenocollema bryospilum grows on Bryopsida

Plant / associate
fruitbody of Rickenella fibula is associated with Bryopsida
Other: sole host/prey

Plant / associate
fruitbody of Rickenella swartzii is associated with Bryopsida
Other: sole host/prey

Foodplant / parasite
fruitbody of Rimbachia arachnoidea parasitises live Bryopsida

Foodplant / parasite
fruitbody of Rimbachia bryophila parasitises live Bryopsida

Foodplant / parasite
fruitbody of Rimbachia neckerae parasitises live Bryopsida

Plant / associate
apothecium of Scutellinia crucipila is associated with Bryopsida
Other: minor host/prey

Plant / grows among
apothecium of Scutellinia paludicola grows among gametophyte of Bryopsida

Plant / grows among
apothecium of Scutellinia trechispora grows among gametophyte of Bryopsida

Foodplant / saprobe
fruitbody of Sistotrema confluens is saprobic on Bryopsida

Foodplant / saprobe
fruitbody of Sistotrema octosporum is saprobic on Bryopsida
Other: unusual host/prey

Plant / grows among
apothecium of Sphaerosporella brunnea grows among gametophyte of Bryopsida
Other: minor host/prey

Plant / associate
fruitbody of Stropharia inuncta is associated with Bryopsida

Plant / associate
fruitbody of Suillus flavidus is associated with Bryopsida

Plant / associate
fruitbody of Tephrocybe confusa is associated with Bryopsida

Plant / associate
fruitbody of Tephrocybe tylicolor is associated with Bryopsida

Plant / associate
fruitbody of Trechispora confinis is associated with live Bryopsida
Other: unusual host/prey

Plant / associate
fruitbody of Trechispora dimitica is associated with live plant of Bryopsida
Other: unusual host/prey

Plant / epiphyte
colony of Trentepohlia aurea grows on Bryopsida

Plant / associate
basidiome of Tulostoma brumale is associated with Bryopsida
Remarks: season: winter - early spring
Other: major host/prey

Plant / associate
basidiome of Tulostoma niveum is associated with Bryopsida
Remarks: season: early winter - early spring

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Evolution and Systematics

Evolution

Discussion of Phylogenetic Relationships

View Bryopsida Tree

Tree from Cox et al (2000), De Luna et al (1999), Goffinet & Cox (2000) and Netwon et al (2000)

The Diphysciales, Funariales, and Timmiales are the most basal lineages within Bryopsida, but these groups have a number of unique features that make comparisons difficult (Netwon et al 2000). The remaining orders are grouped in two large clades: the Dicraniidae consisting of the haplolepidous peristomate orders, and the Bryidae including the diplolepidous-alternate taxa (Figure 3A). The most derived clade within the Bryidae includes orders with ciliate endostomes: Bryales, Rhizogoniales, Hookeriales and Hypnales (Cox et al. 2000, De Luna et al. 1999). The Hedwigiales are interpreted as an instance of peristome loss, and it remains uncertain if they belong in the ciliate clade.

Figure 3. Phylogenetic relationships within the Bryopsida.
Orders in the Dicraniidae in green, orders in the Bryidae in blue.
A. Tree from Newton et al. (2000, p. 195). B. Tree from Goffinet & Cox (2000, p. 218). C. Tree from Cox et al. (2000, p. 235). D. Tree from Cox & Hedderson (1999, p. 126, 132). E. Tree from La Farge et al. (2000, p. 265). F. Tree from De Luna et al (1999, p. 642) and De Luna et al. (2000, p. 251).

As best estimates of higher-level moss phylogeny, recent cladograms of the Bryopsida still contain uncertain relationships for the Funariales, Encalyptales, Bryales, and Hedwigiales.

The Funariales are very likely sister to the rest of arthrodontous mosses (except Diphysiales). This position is consistent in Newton et al. (2000, Figure 3A) and Cox et al. (2000, Figure 3C). However, another analysis places the Funariales as sister to the Encalyptales, and this clade is in turn sister to Dicraniidae (Goffinet & Cox 2000, Figure 3B). The apparent relationship of the Funariales to the Dicraniidae (as interpreted by La Farge et al. 2000, Figure 3E) is most likely because members of the Bryales were used to root the phylogeny. This topology becomes mostly concordant with the other five hypotheses of relationships in Figure 3 if it is rooted with the Funariales.

The relationships of the Encalyptales are still uncertain. This group has been resolved as sister to the Dicraniidae (Newton et al. 2000, Figure 3A), sister to the Bryidae (Cox & Hedderson 1999, Figure 3D), or sister to the Dicraniidae , Bryidae clade (Cox et al. 2000, Figure 3C).

Also the circumscription and relationships of Bryales and Hedwigiales still need further study. Polyphyletic status of the Bryales in several studies make it unclear how families such as the Bartramiaceae, Bryaceae, Mniaceae, and Hedwigiaceae are related to each other.

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records: 7325
Specimens with Sequences: 4527
Specimens with Barcodes: 2137
Species: 2204
Species With Barcodes: 1725
Public Records: 2001
Public Species: 960
Creative Commons Attribution 3.0 (CC BY 3.0)

© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)

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Barcode data

Creative Commons Attribution 3.0 (CC BY 3.0)

© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)

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