Morphology and Classification of Mosses

william r.
buck and bernard goffinet
Morphology and classification of mosses
3.1 Introduction
Bryophytes, in the broad sense, are the second largest phylum of

land plants, after the angiosperms, and inhabit every continent. Among
the bryophytes, mosses (the Bryophyta sensu stricto) are the most spe-
ciose group, comprising approximately 10 000 or more species. They
differ from liverworts and hornworts in a suite of characters (see chapters
1, 2), including macroscopic features such as a gametophyte composed of
stems with undivided and often costate leaves that are typically arranged
all around the axis, and a sporophyte terminated by a capsule that is ele-
vated by the elongation of a seta prior to maturity, and whose mouth is
lined by teeth involved in regulating the dispersal of spores. Mosses
along with liverworts and hornworts represent the oldest lineages
among extant land plants (chapter 5 in this volume). Evidence is mount-
ing that mosses are currently at the highest level of diversity in their evo-
lutionary history (e.g., Kürschner & Parolly 1999). This rather recent
diversification of mosses is likely correlated with the advent of angio-
spermous forests which provide a wide array of habitats. This trend is
best exemplified in tropical rainforests, where most of the diversity in
mosses is found.
Although mosses are rather small organisms, their morphology is rel-
atively complex. Macroscopic characters are limited and most taxonomic
concepts rely on features of the cells composing individual tissues.
Throughout their evolutionary history, mosses have undergone repeated
morphological reduction and simplification (Frey 1981), often as a result
of colonizing specialized, and particularly xeric or ephemeral habitats
(Vitt 1981). Significant innovations or loss thereof occurred early in the
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72 william r. buck and bernard goffinet
diversification of mosses, as revealed by the extreme morphological diffe-

rentiation between the major groups (here recognized as classes, see
below).
The objective of this chapter is to offer a general overview of moss
morphology by describing characters and some of the most widely
encountered states. This brief characterization is followed by a classifica-
tion of all moss genera that we currently recognize.
3.2 Synopsis of moss morphology
Here we present a general overview of the diversity of gametophytic and

sporophytic character-states. For more comprehensive treatments of
moss morphology, the reader is referred to Campbell (1895), Müller
(1895–8), Ruhland (1924), Schofield and Hébant (1984), Vitt (1984), and
Schofield (1985).
3.2.1 Gametophyte
The typical life history of mosses begins with the germination of spores.
The resulting filamentous, uniseriate, branched protonema grows vege-
tatively, and in most taxa is short-lived. (In rare cases germination may be
endosporic, with no obvious protonema.) The protonemata are differen-
tiated into two kinds of filaments, the chloronemata and the caulonemata
(Duckett et al. 1998). The chloronemata are mainly involved in assimila-
tion and propagation (via gemmae). The main role of the caulonema is
colonization. In a few cases (e.g., Sphagnum, Andreaea), the protonemata
are plate-like rather than filamentous (Anderson & Crosby 1965). Gameto-
phores arise laterally from the caulonemata, and it is these leafy, gameto-
phytic stems that are most conspicuous in the moss life cycle. The
gametophyte and sporophyte of mosses are covered by a surface layer at
least analogous to the cuticle of vascular plants, with compounds similar
to those of cuticular waxes of vascular plants (Holloway 1982, Cook &
Graham 1998). Stems may be erect, creeping, or pendent, varying from
less than 1 mm long in mature plants to well over 1 m (e.g., Fontinalis spp.).
Mosses are generally separated into two groups based upon the orienta-
tion and branching of the stems, and the position of gametangia. In acro-
carps (Fig. 3.1) the stems are erect, not or minimally branched, and often
grow in tufts. The apical cell of the stem is eventually used to produce a
terminal gametangium. In some cases growth of the plant is resumed by
the development of subapical innovations which assume the function of
Morphology and classification of mosses 73
Fig. 3.1. Polytrichum piliferum (Polytrichaceae), as an example of an acrocarpous moss.

Note the unbranched, erect stems, sometimes terminating in a sporophyte.
stem. The earliest mosses were probably acrocarpous. For the most part
acrocarpous mosses have colonized terrestrial habitats, especially soil and
rocks. Certainly they are common on the bases of trees, but for a few
notable exceptions, e.g., the Calymperaceae, they have not been success-
ful as epiphytes. The second group of mosses is the pleurocarps (Figs. 3.2,
3.3). The gametangia are produced laterally along the stems and thus the
apical cell continues vegetative growth. These mosses have mostly creep-
ing stems that have extensive lateral branching, resulting in mats. Conse-
quently, virtually all pleurocarpous mosses are perennial, whereas
acrocarps can be either annual or perennial. Although some lineages of
pleurocarps have been successful colonizers of terrestrial habitats (espe-
cially in temperate climates, e.g., Hypnum and Brachythecium), in the
tropics the bulk of the epiphytic mosses are pleurocarps. Growth forms of
mosses have received considerable attention, especially in the ecological
literature. Communities have often been characterized primarily by this
feature alone. The earliest work in this field (Giesenhagen 1910) divided
tropical bryophytes into cushions, turfs, and mats, and individual stems
into unbranched dendroid, branched dendroid, feather, bracket, and
hanging forms. More recent work has divided these various types more
finely (e.g., Mägdefrau 1982). It should be emphasized that growth form is
Fig. 3.2. Helodium blandowii (Helodiaceae), as an example of a pleurocarpous moss.

Note the well-branched stems. Gametangia and sporophytes would be formed
laterally along the stem. Most pleurocarps are prostrate in habit, but this one
grows erect in wet areas.
not necessarily correlated with systematic position (see also La Farge-

England 1996).
Rhizoids
Mosses are typically attached to the substrate by rhizoids. Although
lacking a conductive function, they are not lacking in taxonomic useful-
ness (Koponen 1982). Rhizoids are characterized by being typically
branched, and with diagonal cross-walls between the uniseriately
arranged cells. They may be variously colored or hyaline, smooth or papil-
lose. Rhizoids may arise solely from certain parts of the stems (or rarely
leaves), or they may cover extensive portions of the stems with a felt-like
tomentum. Rhizoids most commonly are of the micronematal type, char-
acterized by a slender diameter, sparse branching, and produced on stems
Fig. 3.3. Brymela websteri (Pilotrichaceae), as an example of a moss. A. Aspect, showing

pleurocarpous habit. B. Short piece of stem with attached branch and
sporophyte, the latter showing seta, capsule, operculum, and calyptra. C.
Young capsule with calyptra. D. Calyptra. E. Operculum. F. Exostome tooth,
showing median furrow. G. Leaves, showing strong double costa, a common
feature of the Hookeriales. H. Leaf apex. I. Laminal cells at mid-leaf. J. Cells at
base of leaf. K. Portion of stem cross-section. L. Axillary hair.
between leaves. In some groups (e.g., the Mniaceae) the rhizoids may be
macronematal, with a larger diameter, extensive branching, and arising
from around branch primordia.
Stems
Stem anatomy has been much used in differentiating taxa at all taxo-
nomic levels (e.g., see Kawai & Ikeda 1970, Kawai 1971–91, Watanabe &
Kawai 1975, Kawai et al. 1985, 1986, Kawai & Ochi 1987, Ron & Kawai 1991).
The external layer, or epidermis, is usually composed of relatively small,

thick-walled cells (as seen in cross-section, Fig. 3.3K), but is sometimes
differentiated as a hyalodermis of enlarged, thin-walled cells. Typically
there is a layer of thick-walled cells subtending the epidermis, providing
structural support for the moss. Internal to this layer is usually a layer
of large, thin-walled parenchyma cells. Although considered “non-
vascular” plants, mosses often have conducting tissue at the center of
their stems. This central strand can be well developed, or completely
lacking. Connections between the conducting tissues of the stem and that
of the leaves occur in some Polytrichales (Hébant 1977). In most other
mosses leaf traces are absent. Hydroids differ from analogous xylem in
vascular plants by lacking lignin (Hébant 1977). Nevertheless, hydroids
share several characteristics with tracheary elements (Schofield & Hébant
1984), which might suggest that they are derived from the same progeni-
tor of vascular tissue.
Stems sometimes are variously ornamented with projections. The
most conspicuous of these structures are paraphyllia. These filamentous
or foliose, simple or branched structures are green and distributed over
most areas of stems (and sometimes branches), especially on older parts.
They are most common in pleurocarpous mosses (e.g., Climacium, Hyloco-
mium, Palustriella). The paraphyllia provide additional photosynthetic
area but their function is otherwise unknown. They may be involved in
external water conductance. Rarely the epidermal cells of stems have
small papillae (e.g., Cyrto-hypnum pygmaeum, Heterocladium macounii). In
stems with branches, there are often other structures which are restricted
to the periphery of branch primordia. These structures are called pseudo-
paraphyllia and may be either filamentous or foliose. Although green,
their function is most probably the protection of branch primordia, not
photosynthetic enhancement. In recent years there has been considerable
work on the origin of pseudoparaphyllia and some authors (Akiyama
1990, Akiyama & Nishimura 1993) have argued that there are two inde-
pendent structures lumped under the single term: true pseudoparaphyl-
lia (almost always filamentous) and “scaly leaves” which are little more
than juvenile leaves. The question deserves a more detailed ontogenetic
study to determine if the various structures originate from the apical cells
of branches (presumably as in scaly leaves) or from the stem surface (pre-
sumably as in true pseudoparaphyllia).
In leaf axils is a structure that has garnered considerable taxonomic
attention in recent years, axillary hairs (Fig. 3.3L). These uniseriate
structures are apparently physiologically active when the leaves are

juvenile (Ligrone 1986) and help keep the expanding leaves from dehy-
drating by secreting a mucilage. They vary in number from one to
several per leaf axil. The basal cell or cells are often differentiated by
being shorter and brownish. The apical cells, varying in number from
one to over ten, are elongate and hyaline. Although often tedious to
observe, they have proved very useful at different taxonomic levels
(Hedenäs 1989b).
The architecture of the gametophyte can be described as modular
(Mishler & De Luna 1991). The hierarchy of the modules is defined by the
mode of branching. Two basic branching patterns occur in mosses:
monopodial and sympodial (La Farge-England 1996). Sympodial
branching occurs in perennial plants with axes of determinate growth.
New branches develop from subapical meristems of the stem or primary
module. These branches grow and function as new stems, sometimes
bearing terminal sex organs. Sympodial branching thus reproduces a
module of identical hierarchical level, and each module is of determi-
nate growth. Sympodial branching is characteristic of all acrocarps and
most epiphytic pleurocarps. In monopodial branching, the develop-
ment of lateral branches is not determined by the cessation of apical
meristematic activity. Lateral meristems develop and grow into
branches, which are often morphologically and functionally distinct
from the stems. Monopodial branching thus involves two distinct
modules, here the stem and the branch. This is the standard pattern seen
in most pleurocarps of the northern hemisphere (e.g., Hypnales sensu
Brotherus 1925). It is typically associated with stable terrestrial habitats
(see During 1979). Some mosses exhibit both modes of branching as they
grow. In the Leucodontales (sensu Brotherus 1925, sub Isobryales) for
example, which are mostly epiphytic, the creeping primary stem turns
away from the substrate and the erect secondary stems often develop
lateral branches. The stem will ultimately stop growing and, at some
distance from the stem apex, a new primary stem arises to resume
growth of the plant. This branching type is the most typical in epiphytic
and to a lesser extent saxicolous mosses. Sympodial branching is likely
the primitive condition in mosses. Monopodial branching is constant
among true pleurocarps, but it has also arisen repeatedly among various
acrocarpous lineages (La Farge-England 1996), and although pleuro-
carps often integrate both modes, they never branch following only a
sympodial pattern.
Leaves
The apical cell of the stem is typically pyramidal with three cutting faces,
but in a few taxa (e.g., Fissidens, Distichium), it has become lenticular and
two-sided, resulting in distichously arranged leaves. In most taxa,
though, phyllotaxy is spiral and leaves are arranged all around the stems
(cf. Berthier 1972). Even in cases where the plants appear strongly flat-
tened, the leaves still arise from all sides of the stems. As in sporophyte-
dominant plants, leaves are the primary photosynthetic organ in the
mosses. Moss leaves are typically formed heteroblastically, with the
basal-most ones on a stem or branch significantly different than those
formed later on the mature stems and branches. For the most part, leaves
vary from lanceolate to ovate (Fig. 3.3G). Unlike in hepatics, moss leaves
are almost never deeply divided into multiple lobes or filaments. The
sole exception is Takakia, in which the leaves are divided into terete fila-
mentous lobes. In some taxa of mosses, virtually all acrocarpous, leaves
may be linear; in other mosses leaves may be orbicular. The leaf surface is
most often flat or slightly concave. However, leaves may also be plicate
(with longitudinal folds), undulate, or rugose. The leaf apex is often
acute to acuminate, but it can also be obtuse, truncate, apiculate, or even
retuse. The leaf insertion is mostly broad, with a wide attachment to the
stem. Sometimes the leaf base may have variously developed auricles and
sometimes the laminal cells on either side of the leaf extend down the
stem as decurrencies. Almost all moss leaves are unistratose. Only in a
few groups are multistratose leaves found, and these are mostly acrocar-
pous (e.g., Grimmiaceae, Diphysciaceae). The largest exception to leaves
being unistratose is marginal thickening. Many mosses, in a large array
of families, have thickened margins (e.g., Fissidens spp., Racomitrium spp.,
Platylomella). In some cases the cells of the margin are differentiated in
shape, color, or wall thickness. Leaves with a border of hyaline, linear
cells are not uncommon. When the margin is differentiated, it is often
referred to as a limbidium. Limbate leaves presumably are an adaptation
resulting in stronger structural integrity. Leaf margins are often not in
the same plane as the rest of the lamina. Commonly the margins are bent
under the leaf and then called recurved. When curled over the leaf, they
are incurved. When the entire leaf is flat, the margins are referred to as
plane. Most moss leaves are toothed on the margins. These teeth may be
entire cells, cell ends, or lateral projections from cells. In almost all cases
the teeth are larger toward the leaf apex, and smaller or absent toward
the leaf base.
Most moss leaves have a costa (or midrib). In the acrocarps, a strong
single costa is predominant. The costa originates from the base of the leaf
(or rarely it is present above but absent below, e.g., some Dicnemonaceae).
A single costa may extend beyond the laminal apex (excurrent), to the
apex (percurrent), or almost anywhere below. The anatomy of the costa
has been extensively used in some groups (especially the Pottiaceae) to
differentiate taxa. Like in the stem, the external layers of the costa are
often thick-walled, but sometimes are enlarged as a hyalodermis, either
on one or both sides of the leaf. Across the middle of the costal cross-
section there is sometimes a row of enlarged guide cells, essentially con-
necting the lamina on either side of the costa. Sometimes there are
extensive areas of slender, very thick-walled cells (sclereids), that may be
either below (abaxial to) the guide cells or on both sides of them. The costa
may be short and double or essentially absent. This costal reduction
seems to have occurred repeatedly in the pleurocarps, but is rare in acro-
carps. The most unusual costal type is the strong double costa (Fig. 3.3G).
This occurs commonly in the Hookeriales but is absent in acrocarps and
rare in the Hypnales.
Some mosses, especially the Polytrichaceae and some Pottiaceae, have
laminal lamellae. These structures, running parallel to the long axis of
the leaf, are mostly unistratose, chlorophyllose sheets of cells. Often the
terminal cell is differentiated from the subtending ones, and such fea-
tures are used in taxonomic recognition. Typically the lamellae are situ-
ated over only the adaxial surface of the costa. Functionally, they are
speculated to primarily increase photosynthetic area. Similar structures
found in some Pottiaceae are uniseriate, chlorophyllose filaments. They
presumably serve a similar function.
Laminal cells may vary in shape from linear to oblate (wider than
long). Most acrocarpous mosses have cells that are relatively short or even
isodiametric. On the other hand, most pleurocarps have elongate laminal
cells (Fig. 3.3H, I). However, most cell shapes can be found in both groups.
Oblate cells in the main part of the lamina are rare, and are seemingly cor-
related with extremely long-linear leaves with a narrow lamina (e.g.,
Calymperes lonchophyllum). The areolation of the Sphagnaceae is unique in
the mosses. In this group, living, chlorophyllose cells (chlorocysts) form a
reticulum that is enclosed by hyaline, hollow cells (hyalocysts). The leaves
are still unistratose, but the green cells may be more exposed on either the
inner or outer surface of the leaf, they may be equally exposed, or they
may be entirely enclosed by the hyaline cells and not exposed on either
surface. An extensive network of pores in the hyalocysts allows rapid

uptake of water into the dead cells. The hyalocysts are typically reinforced
with fibrils. A similar cell dimorphism occurs in the Leucobryaceae and
some Calymperaceae, but here the chlorophyllose cells are embedded
between two layers of leucocysts.
Typically cells are not uniform throughout the moss leaf, in addition
to the possible differentiation of a limbidium. In leaves that have a rela-
tively short apex, i.e., acute to obtuse or truncate, the apical laminal cells
are usually shorter than those at midleaf (e.g., in Platyhypnidium and Neck-
eropsis). In almost all mosses the cells at the leaf insertion are differen-
tiated, typically being longer and sometimes broader (Fig. 3.3J). In many
groups the cells at the basal margins are differentiated. These so-called
alar cells are typically subquadrate. They are sometimes colored or
hyaline, and may be thick-walled or inflated and thin-walled. When there
are leaf decurrencies, sometimes the cells of the decurrencies are similar
to those of the alar regions, and sometimes they are different.
Laminal cells of mosses are sometimes ornamented, and this is more
common in mosses adapted to dry habitats and is thus presumably
adaptational in regard to water relations. The most common type of
ornamentation is papillae. Although grouped under a single term,
papillae are a heterogeneous assemblage of cuticular projections. They
may be solid or hollow, they may be simple, C-shaped, coronate, or vari-
ously branched, and they range from scarcely visible to strongly project-
ing. Cells may have a single papilla, or may have multiple papillae in
various arrangements, and the papillae may be only on one side of the
leaf (most commonly the abaxial surface), or they may be on both sides.
Papillae typically occur over the cell lumina, but sometimes (e.g., Trachy-
pus) are positioned over the cell walls between the lumina. Until fairly
recently another form of cell ornamentation was lumped into papillose,
but is now standardly differentiated. In this case the upper end of the
cell (or less often both ends, or even more rarely just the lower end of the
cell) projects from the surface of the leaf. This is referred to as prorose or
prorulose (depending on the degree of projection). Sometimes laminal
cells are mammillose, with nipple-like projections. In this type of orna-
mentation, the lumen of the cell projects up into the projection, whereas
in papillae the projection is entirely composed of wall material. Rarely
cells have various other kinds of roughenings on their surfaces. When
the upper laminal cells are ornamented, alar cells are almost always
smooth.
Asexual propagula
Mosses have evolved numerous kinds of asexual reproductive structures
(Correns 1899, Newton & Mishler 1994). When water is required for the
transport of sperm to egg, and male and female plants may be quite
distant (sometimes on different continents!), it is no surprise to find effi-
cient mechanisms for vegetative dispersal. Most mosses can grow from
leaf or stem fragments. Typically a leaf or stem fragment will first give rise
to a protonematal stage prior to growth of a new leafy plant. Some taxa
have developed specialized morphological strategies to take advantage of
leaf fragmentation. In these taxa, widely dispersed across the mosses, the
leaf apices are regularly broken off (typically referred to as fragile), either
at the base of a specialized leaf apex (e.g, Groutiella tomentosa), or irregu-
larly along cells lines (e.g., Haplohymenium triste). Sometimes almost the
entire leaf can break into irregular laminal plates (e.g., Zygodon fragilis).
Like in many other groups of plants and fungi, some mosses have devel-
oped a splash-cup mechanism for dispersing small gemmae which, at
maturity, are loose within a cup (e.g., Tetraphis). Some mosses have
evolved specialized gemmae in leaf axils (e.g., Pohlia spp.). These may be
bulbil-like, or more elongate or vermicular; they may be single in leaf
axils or clustered; they may be restricted to the apical portions of the
stems, or throughout. Sometimes elongate, flagellate branches, usually in
dense clusters, are formed in leaf axils and are easily shed (e.g., Platygyr-
ium). These specialized branches have very reduced leaves. Sometimes
specialized stems are produced solely for the production of asexual prop-
agula, with leaves on these stems strongly reduced or absent (e.g., Adelo-
thecium). Asexual reproductive structures are sometimes formed on leaves
(e.g., Calyptopogon, Calymperes, Syrrhopodon). These often are in the form of
few-celled gemmae or filaments. They may arise from specialized cells of
the lamina, either along the margins, at the apex, or throughout, or they
may arise from the costa, either on the abaxial or adaxial surface or both.
Vegetative reproduction is not restricted to just the stems and leaves.
Small bulbils are sometimes formed on rhizoids, presumably as perenni-
ating structures (e.g., Bryum spp.). These may be of different colors and
sizes in different taxa, and thus taxonomically useful (Arts 1989, 1994). In
epiphytic or epiphyllous Hookeriales (esp. Crossomitrium), rhizoidal, fila-
mentous asexual reproductive structures are not uncommon. Asexual
reproductive structures are even known in some groups directly on the
protonemata (e.g., Tetraphidales), but they are not known in the sporo-
phyte generation. The diversity of asexual propagules is matched by a
range of dispersal or abscission mechanisms for these gemmae, as well as

by a variety of germination patterns (Duckett & Ligrone 1992).
Gametangia
The sex organs of mosses are archegonia (female) and antheridia (male).
The flask-shaped archegonium contains a single egg cell in the venter,
which is surmounted by an elongate neck. Numerous sperm cells are pro-
duced in the sack-like antheridium. The arrangement of gametangia in
mosses is varied (Wyatt & Anderson 1984) and is often of taxonomic value.
The two broad groups of sexuality are monoicous (with antheridia and
archegonia on the same plant) and dioicous (with antheridia and archego-
nia on separate plants). In mosses sexuality terms typically have the
ending -oicous whereas in vascular plants they typically end in -oecious,
the latter being strictly applied to sporophytic or diploid sexuality.
Within the monoicous arrangement of gametangia, the most common
variations are autoicous, synoicous, paroicous, and heteroicous (⫽poly-
oicous). In the autoicous arrangement of gametangia the antheridia and
archegonia are in separate inflorescences on the same plant. Within this
broad grouping there is considerable variation, such as cladautoicous
(with the male inflorescence terminal on a separate branch from the
female), gonioautoicous (with the male inflorescence bud-like and axil-
lary on the same stem or branch as the female), and rhizautoicous (with
the male inflorescence on a short branch attached to the female stem by
rhizoids and appearing to be a separate plant). In the synoicous condition
the antheridia and archegonia are mixed within a single inflorescence,
while in paroicous the antheridia and archegonia are in a single inflores-
cence but are not mixed, usually with the sex organs in different leaf axils.
In plants with heteroicous sexuality, there is more than one form of
monoicous inflorescence on the same plant (or species). In one form of
dioicy, the male plants are dwarfed, consisting of little more than the
male inflorescence and a few subtending, reduced leaves, and are epi-
phytic on the female plants, usually either on the leaves or tomentum of
the stem. This is sometimes called a pseudautoicous or phyllodioicous
condition.
In rare cases (e.g., Splachnobryum, Takakia), gametangia are naked in the
axils of vegetative leaves. However, more typically the gametangia are
surrounded and subtended by specialized leaves, usually differentiated
from the vegetative leaves. The archegonia and their subtending leaves
are referred to as perichaetia. The term is typically also used in synoicous
inflorescences. Perichaetial leaves are commonly used taxonomic charac-

ters. They may become either larger or smaller from the outside to the
inside of the inflorescence, and they typically resume growth after fertil-
ization. The antheridia and their subtending leaves are called perigonia.
The perigonial leaves or bracts are usually quite reduced and unlike the
vegetative leaves, especially in autoicous taxa where the perigonia resem-
ble axillary buds. Interspersed among the archegonia and antheridia
within the inflorescences may be sterile hairs, referred to as paraphyses.
Sometimes the apical cell of a paraphysis may be differentiated, e.g., in
Funariaceae.
3.2.2 Sporophyte and associated gametophytic tissues
Seta
The leafless, unbranched sporophyte generation is intimately associated
with the gametophyte. Although green for most of its development, the
sporophyte is physiologically independent only shortly before sporogen-
esis (Paolillo & Bazzaz 1968). The basal-most part of the sporophyte is the
foot, and it is embedded within the gametophyte. At the interface of the
two generations is a transfer zone and the ultrastructural details of this
area (cf. Ligrone & Gambardella 1988) have provided clues about the rela-
tionships of mosses with other land plants (e.g., Ligrone et al. 1993, Frey et
al. 1996). The gametophytic tissue surrounding the foot is referred to as
the vaginula. Traditional wisdom has the vaginula developing from the
basal portions of the archegonium, but considering the occasional occur-
rence of archegonia and hairs on the vaginula, stem tissue seems also to be
involved. The foot is the base of the seta.
The seta is the structure which elevates the sporangium, or spore
capsule, and grows from an apical meristem. In most mosses the seta is
elongate (Fig. 3.3A, B). However, in some reduced taxa (e.g., Acaulon,
Ephemerum), the capsules remain immersed within the perichaetial
leaves because the seta is greatly shortened. The seta is typically smooth,
but in some mosses it is variously roughened (e.g., Brachythecium spp.) or
even spinose (e.g., Calyptrochaeta). It is not uncommon for setae to be
twisted and the pattern and direction of twisting may be taxonomically
useful, e.g., in distinguishing subfamilies of Orthotrichaceae (Goffinet &
Vitt 1998). There is usually a well-developed conducting system within
the seta, even if such tissues are lacking in the gametophyte. Setae are
mostly single per perichaetium, but polysety is known in a few groups
(e.g., Dicranum spp.) where more than one archegonium per perichae-
tium is fertilized. In a few isolated groups (Sphagnum and Andreaea), the
capsule is held aloft from the perichaetial leaves by an extension of
gametophytic tissue, the pseudopodium. The foot of the capsule is
embedded directly in this structure and there is no seta. In Sphagnum, the
capsule is elevated at maturity, rather than long before sporogenesis as in
most mosses.
Capsule
A single capsule develops atop the seta (Fig. 3.3B). In most mosses, cap-
sules are stegocarpous (i.e., regularly dehiscent by shedding an opercu-
lum or lid), but in some groups, typically with immersed capsules, the
capsules are cleistocarpous (i.e., without regular dehiscence), and spores
are dispersed only upon decay and rupture of the capsule wall (Vitt 1981).
Capsule shape varies greatly, from essentially globose to long–cylindric.
Capsules may be symmetric, in which case the capsule is usually either
erect or nutant (i.e., with the apex of the seta sharply bent up to 180° and
the capsule nodding). Alternatively, capsules may be asymmetric. In these
cases, as often found in pleurocarps and some acrocarps, the capsule itself
is bent so that the mouth is oblique to the orientation of the rest of the
capsule. Sometimes capsules are furrowed, either with eight or 16 ribs.
At the base of most capsules there is a differentiated neck, typically cor-
responding to the area below the spore sac within the capsule. In this area
is where one most often finds stomata, although some groups of mosses
lack stomata. The stomata are typically very similar to those in other land
plants, i.e., with two guard cells around a pore. Rarely, there is a single
doughnut-shaped guard cell (e.g., Funaria). In some groups they may be
immersed in small pits (e.g., Orthotrichum spp.). In Sphagnum there are
pairs of guard cells but no pore, and thus the structures are referred to as
pseudostomata (Boudier 1988). In some mosses the neck of the capsule is
greatly exaggerated (e.g., many Splachnaceae) while in others is it little
more than a gradual tapering to the seta. The cells of the capsule wall are
called exothecial cells. In a few groups these may be ornamented with
various projections (e.g., Symphyodon), but for the most part they are
smooth. However, the shape and size of the exothecial cells can be of tax-
onomic value. If a capsule is ribbed, the exothecial cells corresponding to
the area of the ribs are usually narrower and thicker-walled. The cells
directly below the mouth of the capsule are typically different than those
of the rest of the capsule. In most mosses capsules dehisce by the loss of an
operculum (Fig. 3.3B, E). The operculum itself may be almost flat to
long–rostrate. At the mouth of the capsule there are sometimes cells spe-
cifically designed to aid in the dehiscence of the operculum, usually
through swelling with water uptake. These cells, referred to as the
annulus, may not be the result of a single origin. We define the annulus as
differentiated cells which are deciduous, even if tardily so. In many
groups there is a row of cells, usually differentiated in cell wall thickness,
one to several cells high, directly intercalated between the exothecium
and the operculum, which fragment at maturity. This type of annulus is
typical of pleurocarps and some acrocarps. Another type of annulus
found in acrocarps has the differentiated cells inside the capsule, at the
exothecium–operculum junction. These are greatly swollen cells and,
upon dehiscence, the annulus often falls as a complete unit (revoluble
annulus). In an isolated group of mosses, Andreaea, Andreaeobryum, and
Takakia, the capsule dehisces much like in hepatics, by longitudinal slits.
In Andreaea there are typically four (rarely ten), long slits; in Andreaeobryum
there are four to six, short irregular slits; and in Takakia there is a single
spiral slit. It is not clear whether these modes of dehiscence are derived
from a stegocarpous capsule, as suggested for Andreaea by Robinson and
Shaw (1984).
In almost all mosses there is a column of sterile tissue running most of
the length through the capsule. This structure is the columella. It has
received little taxonomic attention but preliminary studies by the senior
author indicate that it may well be a source of valuable systematic charac-
ters, in terms of shape (including cross-sectional) and length, especially in
the acrocarps. In a few groups the apex of the columella is fused to the
operculum (e.g., Schistidium). In this systylius condition, the columella
eventually breaks free from its connection with the apex of the seta and
falls with the operculum. In the Polytrichaceae the apex of the columella
is expanded into a circular membrane, the epiphragm, which partially
closes the capsule mouth after dehiscence. In Sphagnum, Andreaea, and
Andreaobryum the columella is short, and the spore sac, rather than being
cylindrical, is dome-shaped. Only in Archidium is a columella completely
missing (Snider 1975).
Peristome
The capsule mouth of most stegocarpous mosses is lined by teeth, which
together form the peristome (Fig. 3.3F). The peristome offers a variety of
characters, some of which, such as the architecture and the pattern of cell
division in particular, are central to most modern higher-level classifica-

tions of mosses (e.g., Dixon 1932, Vitt 1984, Vitt et al. 1998). Much work has
been done in recent years on peristome architecture (e.g., Edwards 1979,
1984) and development (e.g., Shaw et al. 1987, 1989a, b, Shaw & Anderson
1988). Only a brief overview of the subject can be presented here. As in the
gametophyte, growth of the sporophyte is insured by the meristematic
activity of an apical cell. In the early stages of development (cf. Wenderoth
1931, Snider 1975, for a complete sequence of zygotic divisions) each cell
derived from the apical cell immediately undergoes an anticlinal divi-
sion. By the time two derivatives have been produced and each has under-
gone a division, four cells are seen in transverse section below the apical
cell. Subsequent divisions lead to the differentiation of two fundamental
tissues, the endothecium and the amphithecium (Wenderoth 1931, Shaw
et al. 1987, 1989a, b). This organization is known as the fundamental
square (Goebel 1905). Each of the four merophytes will follow an identical
pattern of cell divisions. The endothecium will give rise to the columella
and sporogenous tissue in most mosses, except Sphagnum where the spor-
ogenous tissue is of amphithecial origin. From the amphithecium are
derived the exothecial cells and their underlying parenchyma, which in
the apical portion of the capsule are differentiated into the peristome
forming layers. These innermost three layers of the amphithecium are
called the outer, the primary, and the inner peristomial layers (OPL, PPL,
and IPL, respectively), with the latter being adjacent to the endothecium
(Blomquist & Robertson 1941).
Edwards (1979) proposed the peristomial formula to describe the
numbers of cells in each of the three peristomial layers contributing to
the peristome. For ease of comparison, the peristomial formula uses the
number of cells in 1 ⁄ 8 of the capsule circumference. For example, a 4:2:4
formula means that in a whole capsule there are 32, 16, and 32 cells in the
OPL, PPL, and IPL, respectively. Haplolepideous peristomes have a
formula of 4:2:3 and diplolepideous peristomes have formulae ranging
from 4:2:4 to 4:2:14. The peristome of Funaria has a formula of 4:2:4 (Fig.
3.4A, C). In addition, the peristome of Funaria has endostome segments
opposite the exostome teeth and all anticlinal walls in the IPL are aligned
with anticlinal walls of the PPL. In most other diplolepideous mosses,
where the endostome segments alternate with the exostome teeth, anti-
clinal walls of IPL cells are offset with regard to those in the PPL. Thus, the
4:2:4 formula of Funaria develops somewhat differently than the 4:2:4
formula of a Pohlia (Fig. 3.4D), for example.
Fig. 3.4. Architecture of arthrodontous peristomes at maturity. One-eighth of the

amphithecium is represented, with the corresponding peristomial formula
indicating the number of cells found in one-eighth of each layer. Cell-wall
thickening is represented in heavy black areas; fine lines correspond to walls
that will be almost completely resorbed in the latest stages of development.
A. Diplolepideous opposite peristome of Funaria. B. Haplolepideous peristome;
note that each segment of the endostome is composed on its inner surface of
one and a half cells. C. Diplolepideous alternate peristome as typically found
in the Orthotrichaceae; for every two segments the inner peristomial layer
(IPL) is composed of four cells at most, the inner surface of the segments bears
a vertical line, although it is not median, or rarely so, depending on the degree
of displacement of anticlinal walls during maturation, the outer peristomial
layer (OPL) is thicker than the primary peristomial layer (PPL), and the
exostome teeth are found paired rather than free. D. Typical diplolepideous
alternate peristome (e.g., in the Bryidae and Hypnidae); note that the IPL has
undergone additional divisions, leading to six cells per pair of segments, and
to the development of cilia in between the segments; the inner surface of the
segments never bears a median line, as all anticlinal IPL walls are displaced
during maturation.
During the final stages of sporophyte maturation, additional material

is deposited on the cell walls of the inner layers of the amphithecium. In
the Polytrichopsida at maturity each tooth is composed of several elon-
gate whole cells, and the peristome is designated as nematodontous (Fig.
3.5). In most other peristomate mosses, the cells are short and wide, and
only periclinal walls are thickened while longitudinal anticlinal walls are
resorbed so that at maturity the amphithecial layers appear as rows of
teeth composed of cell wall remnants, i.e., the arthrodontous peristome
(Figs. 3.6–3.8). Each tooth is composed of periclinal walls from two adja-
cent amphithecial cells, separated by their middle lamellae. The degree of
thickening may vary between layers, thereby accounting for the hygro-
scopic nature of the peristome or how the peristome responds to changes
in ambient moisture: a heavier outer thickening results in the teeth
bending outward as they dry, while a thicker inner surfaces makes the
peristome bend inward.
Arthrodontous peristomes can be classified into two general types:
Fig. 3.5. Tetraphis pellucida (Tetraphidaceae). Light micrograph of two of the four
peristome teeth in the nematodontous peristome.
diplolepideous (Fig. 3.8) and haplolepideous (Figs. 3.6, 3.7). In the former,
two concentric rows of teeth are developed around the capsule mouth,
and the outer surface of the outer teeth bears a median line marking the
junction of two rows of cells. By contrast, the outer surface of the haplole-
pideous tooth lacks such a line because it is made from a single column of
cells. Although the diplolepideous peristome is typically composed of
Fig. 3.6. Schistidium heterophyllum (Grimmiaceae). Light micrograph of portion of the

haplolepideous peristome.
two rows of teeth, an exostome and an endostome, while haplolepideous

taxa typically bear only a single row (homologous to the endostome of the
Diplolepideae), the terms refer solely to the number of cells composing
the outer surface of the teeth. Since exostomes are lacking or strongly
reduced in the Haplolepideae, it is clear that the terms compare two non-
homologous characters, namely the outer surface of the exostome of the
Diplolepideae with that of the endostome of the Haplolepideae.
The exostome is composed of 16 teeth. These can each be divided in
Fig. 3.7. Tortula plinthobia (Pottiaceae). SEM of portion of the haplolepideous

peristome.
two or they can be fused into eight pairs. The thickening of the surfaces
of the teeth or segments often follows a pattern that is characteristic for
the family, genus or, more rarely, species. The inner surface of the exos-
tome is typically not very ornamented, but commonly there are striking
projections associated with the cell walls; this condition is referred to as
trabeculate. In its most developed condition, the endostome of a diplo-
lepideous taxon has a tall basal membrane, essentially a tube of tissue.
Perched upon the basal membrane are 16 segments, most often alternat-
ing in position with the exostome teeth. These segments are often
keeled and perforate and extend to about the same level as the apex of
the exostome teeth. Alternating on the basal membrane with the seg-
ments, and thus typically opposite the exostome teeth, are groups of
cilia. The presence of cilia is correlated with additional divisions in the
IPL; they are always lacking in the Funariales and the Orthotrichales
which consistently have 32 or fewer IPL cells, and present in many
Bryales with 48 or more cells in the IPL. The cilia are mostly uniseriate
structures and occur in groups of one to three or four. In some taxa, cilia
bear attachments (then referred to as appendiculate) or are swollen at the
Fig. 3.8. Pseudoscleropodium purum (Brachytheciaceae). SEM of portion of the

diplolepideous peristome, showing median line on front surface of exostome
and projecting trabeculae on the back surface, and an endostome with a high
basal membrane, alternating, perforate segments, and cilia in groups of three.
nodes or joints between two cells of a column (nodulose). The peristome

of haplolepideous taxa may have individual, free teeth, sometimes
divided almost to their base into filaments, or sometimes the teeth are
fused into an extensive tube, with or without free filaments at its apex.
When selective pressures on the peristome are relaxed, or if selection
favors a reduced peristome, the endostome tends to be reduced prior to
the exostome.
When the capsule is young and not fully developed, there is a structure
of gametophytic tissue which encloses it. Near capsule maturity this struc-
ture, the calyptra, falls and frees the capsule, allowing dehiscence (Fig.
3.3B–D). In a few cases it stays attached to the apex of the seta. The calyptra
is formed from the expanded upper portions of the archegonium. There
are two basic forms of calyptrae, mitrate and cucullate. The mitrate calyp-
tra is conic and undivided (similar to a bishop’s miter) or equally lobed at
the base. A cucullate calyptra is split up one side, and is similar to a hood,
being longer on one side than the other. There seems to be a correlation
between obliquely rostrate opercula and cucullate calyptrae, and between

erect–rostrate opercula and mitrate calyptrae. Calyptrae can be naked or
hairy, smooth or roughened. Because of the limited variability found in
calyptrae, convergent evolution has occurred repeatedly.
Spores
Moss spores are typically single-celled, spherical, and about 12–15 ␮m in
diameter. In some cases the spores may be multicellular (e.g., Dicnemona-
ceae), typically associated with endosporic germination and relatively
xeric environmental conditions. Some spores are bilaterally symmetrical
or retain tetrad markings (or rarely remain in tetrads at maturity). Spores
range in size from about 5 ␮m to well over 100 ␮m. For the most part moss
spores are finely papillose, but range from essentially smooth to highly
ridged, spinose, or reticulate. Strongly ornamented spores are often large
and associated with cleistocarpous capsules. Spore coats are multilayered;
their structure was reviewed by Mogensen (1983). Spore number per
capsule is typically in the hundreds of thousands, but at the opposite
extreme it may be as few as four (in Archidium).
In most stegocarpous mosses a peristome is attached around the
circumference of the capsule mouth and is associated with spore disper-
sal. The peristome is typically attached to the spore sac and, when the peri-
stome moves back and forth through hygroscopic movement, the spore
sac is moved, keeping the spores from becoming compacted in the base of
the capsule (Pfaehler 1904). However, the peristome has additional func-
tions in spore dispersal as well. Through hygroscopic motion, the peri-
stome either seals the mouth of the capsule or opens it. Some mosses open
their capsule mouths under dry conditions (xerocastique), and others
open their mouths when moist conditions prevail (hygrocastique)
(Mueller & Neumann 1988). In diplolepideous mosses the endostome is
not conspicuously hygroscopic, but in the Haplolepideae the hygroscopic
function characteristic of the diplolepideous exostome is carried out by
its peristome, the endostomial homolog (Crum 1977). Additionally, peri-
stomes often act as filters, allowing spores to be released over extended
periods of time. Peristomes may even be active spore dispersal agents by
having spores get caught in the exostomial trabeculae and, through
hygroscopic movement, flick the spores away from the capsule. Spore dis-
persal can be further enhanced by modifications of the capsule wall.
Alternating bands of thick-walled and thin-walled exothecial cells result
in the capsule contracting when dry, and thereby pushing spores up. In
the Splachnaceae, differential thickening of walls is within individual

cells. Here the transverse walls are thicker than the longitudinal ones and
as a result, the capsule shrinks when it dries and the spore mass is pushed
upward (Koponen 1990). In Tetraplodon the efficiency of this mechanism is
improved by the presence of a pseudocolumella, which is located within
the neck (or hypophysis) and pushes the whole spore sac up (Demidova &
Filin 1994). Capsule contraction is also the mechanism by which Scouleria
ejects its spores (Churchill 1985). In Sphagnum, the capsule wall bears
pseudostomata which are arranged in a more or less median band and
allow the capsule to contract quickly (Boudier 1988) leading to the dehis-
cence of the operculum and explosive spore dispersal. In valvate capsules,
such as in Andreaea, the valves bend outward upon drying, thereby expos-
ing the spore mass. Despite these various innovations, spore dispersal in
most mosses is rather passive, relying on wind as the dispersing agent.
The only exception is found in the Splachnaceae, where insects are the
primary dispersal agent (Marino 1997). Such a dispersal mode is referred
to entomophily (or more appropriately entomochory) and works by
deceiving insects. Insects looking for fresh dung or cadavers on which to
lay their eggs are attracted to the capsules of these mosses which emit vol-
atile compounds through their stomata, the odor of which is reminiscent
of decaying animal remains or dung. The deceived insect subsequently
carries the sticky spores to a true nitrogenous substrate, upon which the
spores germinate.
3.3 The classification of mosses
Throughout the lifetimes of most living bryologists classification systems

have been proposed to reflect natural relationships. Of course through
time our understanding of phylogeny and evolutionary processes has
greatly changed. Nevertheless, through most of recent time, bryologists
have attempted to understand how mosses are related, and then used
this knowledge to construct classifications. Since the publication of
Brotherus’ treatment of the mosses in the second edition of Engler’s Die
natürlichen Pflanzenfamilien in 1924–5, moss classification has been greatly
swayed by peristomial morphology and ornamentation. During prepara-
tion of this work, Brotherus was heavily influenced by Fleischer’s monu-
mental Die Musci der Flora von Buitenzorg (1904–23). Fleischer himself was
influenced by the work of Philibert (1884–1902), who pioneered studies on
peristome structure. Consequently, for most of the current century,
details of the peristome were considered as valuable in understanding

relationships as were flowers within the angiosperms. However, as noted
above, peristomes are vital structures in spore dispersal and thus are as
subject to environmental pressures as are any other part of the moss
plant, and should therefore not be regarded of greater value for higher-
level classification than gametophytic characters (Buck 1991). It has long
been documented that the epiphytic condition is correlated with reduc-
tions in sporophytic characters (Mitten 1859). Capsules often become
erect, peristomes become inserted below the capsule mouth, endostomial
cilia are lost, the basal membrane becomes lower, eventually the endo-
stome may be lost entirely, the inner surface of the exostome becomes less
trabeculate, the ornamentation on the outer surface of the exostome
becomes less, and in some extreme circumstances, the exostome is also
lost. Nevertheless, there are certainly peristomial characters that remain
valuable and diagnostic as taxonomic markers. In the diagnoses asso-
ciated with the following classification, peristome characters are fre-
quently used, and indeed seem to be extremely valuable. It is just the
unquestioning use of peristome characters in the face of overwhelming
contradictory evidence that should be challenged.
Among peristomial characters that are used in classification is the
standard dichotomy of haplolepideous and diplolepideous. However, it
appears as if the haplolepideous mosses arose from within the diplolepid-
eous mosses and thus recognition of a formal grouping for all haplolepid-
eous mosses leaves the diplolepideous mosses paraphyletic. Nevertheless,
the haplolepideous mosses, on this single peristome character, are a
natural group. Peristomial ornamentation has long been used for classifi-
cation. For example, teeth with vertical pits characterize the Dicranaceae.
Within the pleurocarps peristomial ornamentation has less value because
of the repeated invasion by various lineages into epiphytic habitats with
consequent sporophytic reductions. Thus the terrestrial taxa, most of
which have striking horizontal striations on the external surface of the
exostome, do not form a natural group, but rather an ecological one.
Therefore, suites of correlated characters are needed to arrive at a substan-
tiative classification.
Because of its more complex morphology, the gametophyte genera-
tion offers a bonanza of characters that can be used in classification
(Hedenäs 1989a). Without going into great detail, characters that have
been used for understanding relationships can be found thoughout the
gametophore. These range from details of stem anatomy, to rhizoidal
morphology and insertion, to structure of pseudoparaphyllia, to

morphology of axillary hairs. These have been correlated with tradition-
ally used gametophytic characters of leaf areolation and costal develop-
ment.
Certainly there are still morphological characters that offer great
promise in helping to better understand relationships that are currently
not being used. Examples include columellar morphology and stem epi-
dermal patterns in relationship to leaf insertion. Surely more will surface
with increased observation. In recent years, though, molecular characters
are having more influence on classifications than any other set of charac-
ters. Although these can be very powerful data sets, caution is needed.
Single-gene classification systems are just as likely to be faulty as are any
other type of single-character classification. This is easily observable
when sequence data from different genetic loci reveal different phyloge-
nies. They cannot all be correct. Evolution of a genetic locus or a plastid is
not identical to that of the whole organism. Nevertheless, the use of
sequence data cannot and should not be ignored. In groups of organisms
with frequent morphological reduction, such as the mosses, insights
from molecular data allows us to re-evaluate morphological data
(Goffinet et al. 1998, Buck et al. in press).
3.4 A suggested classification of the mosses
The following classification is original in several aspects. Molecular data

have been very influential (e.g., Goffinet et al. 1998, Cox & Hedderson
1999, Beckert et al. 1999, Buck et al. in press), but in the many cases where
such data are lacking, morphology has been used exclusively. In some
cases the morphological data have been interpreted differently than in
the past. To date there are no cladistic analyses of morphological charac-
ters for mosses addressing the entire spectrum of families and orders.
Undoubtedly, this classification, which is little more than a working
hypothesis, will be a target. New information with implications for phy-
logeny and classification is being uncovered at a rapid pace. Thus,
although this classification is our best estimate at the time of writing
(September 1999), we are certain that in short order our own ideas will
change. This is the nature of progress. The best that we can hope for is
that some of our novel ideas will prove accurate and that the others will
encourage bryologists to become more involved in this kind of basic
systematic research. Please note that within the subclass Bryidae we have
adopted superorders, following Reveal’s (1998) suggestion to use the

ending -anae, even though Reveal’s specific proposal was defeated during
the nomenclature session at the St Louis International Botanical Con-
gress.
PHYLUM: BRYOPHYTA
CLASS: TAKAKIOPSIDA Leaves divided into terete filaments; capsules

dehiscent by a single longitudinal spiral slit.
order: Takakiales
Takakiaceae S. Hatt. & Inoue. Takakia S. Hatt. & Inoue
CLASS: SPHAGNOPSIDA Branches usually in fascicles; leaves
composed of a network of chlorophyllose and hyaline cells; setae lacking;
capsules elevated on a pseudopodium; stomata lacking.
order: sphagnales Plants mostly branched, with branches in fascicles;
stems with wood cylinder; leaves unistratose; antheridia subglobose;
archegonia terminal on branches; capsules ovoid.
Sphagnaceae Dum. Sphagnum L.
order: ambuchananiales Plants sparsely branched, with branches not in
fascicles; stems without wood cylinder; leaves partially bistratose; antheridia
oblong–cylindric; archegonia terminal on stems; capsules cylindrical.
Ambuchananiaceae Seppelt & H. A. Crum. Ambuchanania Seppelt &
H. A. Crum
CLASS: ANDREAEOPSIDA Plants on acidic rocks, generally autoicous;
cauline central strand absent; calyptrae small; capsules valvate, with four
valves attached at apex; seta absent, pseudopodium present.
order: andreaeales
Andreaeaceae Dum. Acroschisma Lindl., Andreaea Hedw.
CLASS: ANDREAEOBRYOPSIDA Plants on calcareous rocks,
dioicous; cauline central strand lacking; calyptrae large and covering
whole capsule; capsules valvate, apex eroding and valves free when old;
seta present.
order: andreaeobryales
Andreaeobryaceae Steere & B. M. Murray. Andreaeobryum Steere &
B. M. Murray
CLASS: POLYTRICHOPSIDA Plants typically robust, dioicous; cauline
central strand present; leaves costate, often with ventral lamellae on
costa; capsules operculate; seta present; peristome nematodontous.
order: Tetraphidales Gametophore relatively small; stems not
rhizomatous; costa slender, elamellate; capsules long–exserted.
Tetraphidaceae Schimp. Leaves unicostate; calyptrae small conic;

capsule symmetric and erect, neck short; peristome single, of four
erect teeth; Tetraphis Hedw. (Fig. 5), Tetrodontium Schwägr.
Oedipodiaceae Schimp. Leaves unicostate; calyptrae cucullate; capsule
symmetric and erect, neck very long; capsules gymnostomous;
Oedipodium Schwägr.
Buxbaumiaceae Schimp. Leaves ecostate; calyptrae cucullate or
mitrate; capsule strongly asymmetric and horizontal, neck short;
peristome double; Buxbaumia Hedw.
order: polytrichales Gametophore typically large; stems typically
rhizomatous; costa broad, with adaxial chlorophyllose lamellae; peristome
mostly of (16)32–64 teeth.
Polytrichaceae Schwägr. Alophosia Card., Atrichopsis Card., Atrichum P.
Beauv., Bartramiopsis Kindb., Dawsonia R. Br., Dendroligotrichum (Müll.
Hal.) Broth., Hebantia G. L. S. Merr., Itatiella G. L. Sm., Lyellia R. Br.,
Meiotrichum (G. L. Sm.) G. L. S. Merr., Notoligotrichum G. L. Sm.,
Oligotrichum Lam. & DC., Plagioracelopus G. L. S. Merr., Pogonatum P.
Beauv., Polytrichadelphus (Müll. Hal.) Mitt., Polytrichastrum G. L. Sm.,
Polytrichum Hedw. (Fig. 1), Pseudatrichum Reimers, Pseudoracelopus
Broth., Psilopilum Brid., Racelopodopsis Thér., Racelopus Dozy & Molk.,
Steereobryon G. L. Sm.
CLASS: BRYOPSIDA Plants small to robust; leaves costate, typically
lacking lamellae; capsules operculate; peristome arthrodontous.
subclass: diphysciidae Gametophore small, perennial; leaves
costate; capsules immersed among long perichaetial leaves; peristome
double.
order: diphysciales Plants perennial; leaves costate; capsules immersed
among long perichaetial leaves.
Diphysciaceae M. Fleisch. Diphyscium D. Mohr, Muscoflorschuetzia
Crosby, Theriotia Cardot
subclass: funariidae Plants terricolous, acrocarpous; stem typically
with central strand; annulus often well developed.
order: timmiales Plants robust; laminal cells mammillose; calyptrae
cuculate, often persistent; capsules bent to pendent; cilia present.
Timmiaceae Schimp. Timmia Hedw.
order: encalyptales Plants small to medium-size; laminal cells thick-
walled, isodiametric above, rectangular and hyaline or reddish below; annulus
not differentiated; calyptrae very large, enclosing the entire erect capsule.
Encalyptaceae Schimp. Bryobrittonia R. S. Williams, Encalypta Hedw.
order: funariales Peristome lacking cilia, opposite, divisions in the IPL

symmetric.
Funariaceae Schwägr. Protonema short-lived; costa well developed;
laminal cells smooth and thin-walled; paraphyses with swollen
apical cell; calyptrae smooth and naked; stomata with single guard
cell; peristomes opposite, following a 4:2:4 pattern, or lacking;
Aphanorrhegma Sull., Brachymeniopsis Broth., Bryobeckettia Fife,
Corynotheca Ochyra, Cygnicollum Fife & Magill, Entosthodon Schwägr.,
Funaria Hedw., Funariella Sérgio, ⫻Funariophyscomitrella Wettst.,
Loiseaubryum Bizot, Nanomitriella E. B. Bartram, Physcomitrella Bruch &
Schimp., Physcomitrellopsis Broth. & Wager, Physcomitrium (Brid.) Brid.,
Pyramidula Brid.
Disceliaceae Schimp. Protonemata persistent; costa weak to absent;
calyptrae persistent below the urn; stomata lacking; peristomes
reduced and opposite; Discelium Brid.
Gigaspermaceae Lindb. Gametophore united by subterranean
rhizomatous aphyllous stem, cladocarpous; annulus not
differentiated; capsules often immersed, gymnostomous;
Chamaebryum Thér. & Dixon, Costesia Thér., Gigaspermum Lindb.,
Lorentziella Müll. Hal., Neosharpiella H. Rob. & Delgad., Oedipodiella
Dixon
subclass: dicranidae Peristome haplolepideous, with a formula of
(4):2:3; exostome typically absent; late state division in the IPL
asymmetric.
order: grimmiales Plants slender to robust; laminal cells with thick and
often wavy walls; peristome of 16 entire or divided teeth.
Grimmiaceae Arn. Laminal cells with sinuose walls; leaves often
terminated by hair-point; Aligrimmia R. S. Williams, Coscinodon
Spreng., Coscinodontella R. S. Williams, Dryptodon Brid., Grimmia
Hedw., Indusiella Broth. & Müll. Hal., Jaffueliobryum Thér.,
Leucoperichaetium Magill, Racomitrium Brid., Schistidium Bruch &
Schimp. (Fig. 6)
Ptychomitriaceae Schimp. Laminal cells with straight walls; leaves
lacking hair-points; Campylostelium Bruch & Schimp., Glyphomitrium
Brid., Ptychomitriopsis Dixon, Ptychomitrium Fürnr.
Scouleriaceae S. P. Churchill. Plants blackish, acro- or cladocarpous,
saxicolous in riparian habitats; calyptrae mitrate, smooth; annulus
not differentiated; capsules urceolate to globose; Scouleria Hook.,
Tridontium Hook. f.
Drummondiaceae (Vitt) Goffinet, comb. nov. (Orthotrichaceae

subfam. Drummondioideae Vitt, Canad. J. Bot. 50: 1194. 1972). Stem
with central strand, cladocarpous; costa with differentiated adaxial
stereids; laminal cells thick-walled; peristome reduced; Drummondia
Hook.
order: archidiales Plants small, ephemeral; seta lacking; capsules
cleistocarpous, with less than 200 large spores (often 4–8); columella lacking.
Archidiaceae Schimp. Archidium Brid.
order: seligeriales Plants generally saxicolous; alar cells differentiated;
capsules often systylious.
Seligeriaceae Schimp. Plants small; leaves costate; costa homogenous
in transverse section; peristome mostly deeply inserted, relatively
well developed; spores small; Blindia Bruch & Schimp., Brachydontium
Fürnr., Hymenolomopsis Thér., Seligeria Bruch & Schimp., Trochobryum
Breidl. & Beck
Wardiaceae W. H. Welch. Plants robust, rheophilic; leaves ecostate;
peristome attached at mouth, reduced; spores large; Wardia Harv. &
Hook.
order: dicranales Plants small to large; laminal cells generally smooth;
alar cells often differentiated; peristome single, lacking basal membrane,
segments trabeculate and striate.
Bryoxiphiaceae Besch. Leaves distichous with small dorsal extension
(?) along costa; capsules gymnostomous; Bryoxiphium Mitt.
Fissidentaceae Schimp. Leaves distichous and complanate, with
vaginant lamellae; apical cell two-sided; Fissidens Hedw.
Dicranaceae Schimp. Plants generally robust; cauline central strand
present; leaves with well differentiated alar cells; peristome of 16 flat
teeth divided in upper two-thirds, typically with vertically pitted
outer surface; Anisothecium Mitt., Aongstroemia Bruch & Schimp.,
Aongstroemiopsis M. Fleisch., Arctoa Bruch & Schimp., Atractylocarpus
Mitt., Braunfelsia Paris, Brothera Müll. Hal., Brotherobryum M. Fleisch.,
Bryohumbertia P. de la Varde & Thér., Bryotestua Thér. & P. de la Varde,
Camptodontium Dusén, Campylopodiella Cardot, Campylopodium (Müll.
Hal.) Besch., Campylopus Brid., Chorisodontium (Mitt.) Broth., Cnestrum
I. Hagen, Cryptodicranum E. B. Bartram, Dicranella (Müll. Hal.)
Schimp., Dicranodontium Bruch & Schimp., Dicranoloma (Renauld)
Renauld, Dicranum Hedw., Eucamptodontopsis Broth., Holomitriopsis H.
Rob., Holomitrium Brid., Hygrodicranum Cardot, Kiaeria I. Hagen,
Kingiobryum H. Rob., Leptotrichella (Müll. Hal.) Lindb., Leucoloma Brid.,
Macrodictyum (Broth.) E. H. Hegew., Mesotus Mitt., Microcampylopus

(Müll. Hal.) Fleisch., Mitrobryum H. Rob., Muscoherzogia Ochyra,
Orthodicranum (Bruch & Schimp.) Loeske, Paraleucobryum (Limpr.)
Loeske, Parisia Broth., Pilopogon Brid., Platyneuron (Cardot) Broth.,
Pocsiella Bizot, Polymerodon Herzog, Pseudephemerum (Lindb.) I. Hagen,
Pseudochorisodontium (Broth.) C. H. Gao, Vitt, X. H. Fu & T. Cao,
Schliephackea Müll. Hal., Sclerodontium Schwägr., Sphaerothecium
Hampe, Steyermarkiella H. Rob., Symblepharis Mont., Werneriobryum
Herzog
Bruchiaceae Schimp. Alar cells not differentiated; capsules with
elongate necks; spores mostly with trilete markings, usually strongly
ornamented; Bruchia Schwägr., Cladophascum Sim, Eobruchia W. R.
Buck, Pringleella Cardot, Trematodon Michx.
Dicnemonaceae Broth. Plants robust, cladocarpous; central strand
generally absent; laminal cells elongate, thick-walled and porose;
perichaetial leaves differentiated and sheathing; calyptrae mitrate;
germination endosporous; Dicnemon Schwägr., Eucamptodon Mont.
Leucobryaceae Schimp. Plants robust, glaucous; cauline central strand
lacking; costa broad, occupying most of the leaf, with median
chlorophyllose cells and adaxial and abaxial layers of hyaline cells;
Cladopodanthus Dozy & Molk., Leucobryum Hampe, Ochrobryum Mitt.,
Schistomitrium Dozy & Molk.
Ditrichaceae Limpr. Plants slender; alar cells not differentiated;
peristome of 16 completely divided, terete teeth; Astomiopsis Müll.
Hal., Austrophilibertiella Ochyra, Bryomanginia Thér., Ceratodon Brid.,
Cheilothela Broth., Chrysoblastella R. S. Williams, Cladastomum Müll.
Hal., Cleistocarpidium Ochyra & Bednarek-Ochyra, Crumuscus W. R.
Buck & Snider, Cygniella H. A. Crum, Distichium Bruch & Schimp.,
Ditrichopsis Broth., Ditrichum Hampe, Eccremidium Hook. f. & Wilson,
Garckea Müll. Hal., Kleioweisiopsis Dixon, ⫻Pleuriditrichum A. L.
Andrews & F. J. Herm., Pleuridium Rabenh., Rhamphidium Mitt.,
Saelania Lindb., Skottsbergia Cardot, Strombulidens W. R. Buck, Trichodon
Schimp., Tristichium Müll. Hal., Wilsoniella Müll. Hal.
Rhabdoweisiaceae Limpr. Plants small to medium size; stem lacking
central strand; capsules ribbed, widest at mouth; Amphidium Schimp.,
Cynodontium Schimp., Dichodontium Schimp., Dicranoweisia Milde,
Holodontium (Mitt.) Broth., Oncophorus (Brid.) Brid., Oreas Brid.,
Oreoweisia (Bruch & Schimp.) De Not., Pseudohyophila Hilp.,
Rhabdoweisia Bruch & Schimp., Verrucidens Cardot
Rhachitheciaceae H. Rob. Laminal cells rectangular in lower half,

short to isodiametric above; alar cells not differentiated; perichaetial
leaves differentiated; capsules ribbed, rarely smooth; endostome
teeth fused or not; IPL of 8 or 16 cells only (peristome formula: (4):2:2
or (4):2:1); Hypnodontopsis Z. Iwats. & Nog., Jonesiobryum B. H. Allen &
Pursell, Rhachitheciopsis P. de la Varde, Rhachithecium Jolis, Tisserantiella
P. de la Varde, Uleastrum W. R. Buck, Zanderia Goffinet
Erpodiaceae Broth. Plants cladocarpous; costa lacking; laminal cells
often papillose; calyptrae mitrate; Aulacopilum Wilson, Bryowijkia
Nog., Erpodium (Brid.) Brid., Solmsiella Müll. Hal., Venturiella Müll.
Hal., Wildia Müll. Hal. & Broth.
Schistostegaceae Schimp. Gametophores dimorphic, small, annual,
arising from persistent luminescent protonemata; leaves ecostate,
distichous or in five rows; capsules globose, gymnostomous, lacking
stomata and annulus; Schistostega D. Mohr
Eustichiaceae Broth. Leaves distichous; laminal cells quadrate and
thick-walled; capsules ribbed; peristome of 16 teeth; Eustichia (Brid.)
Brid.
Viridivelleraceae I. G. Stone. Protonemata persistent; stems producing
gametangia and associated leaves only; capsules gymnostomous;
Viridivellus I. G. Stone
order: pottiales Plants minute to robust, generally orthotropic; upper
laminal cells usually isodiametric and papillose; alar cells not differentiated;
perichaetial leaves typically not differentiated; capsules erect; peristome
typically papillose, not trabeculate.
Pottiaceae Schimp. Plants small to robust, primarily terrestrial; cauline
central strand often present; leaves narrowly lanceolate to ligulate;
laminal cells typically papillose; calyptrae cucullate, naked, smooth;
peristome of 16 or 32 segments; Acaulon Müll. Hal., Aloinella Cardot,
Aloinia Kindb., Anoectangium Schwägr., Aschisma Lindb., Barbula Hedw.,
Bellibarbula P. C. Chen, Bryoceuthospora H. A. Crum & L. E. Anderson,
Bryoerythrophyllum P. C. Chen, Calymperastrum I. G. Stone, Calyptopogon
(Mitt.) Broth., Chenia R. H. Zander, Chionoloma Dixon, Crossidium Jur.,
Crumia W. B. Schofield, Dialytrichia (Schimp.) Limpr., Didymodon Hedw.,
Dolotortula R. H. Zander, Erythrophyllastrum R. H. Zander,
Erythrophyllopsis Broth., Eucladium Bruch & Schimp., Ganguleea R. H.
Zander, Gertrudiella Broth., Globulinella Steere, Goniomitrium Hook.f. &
Wilson, Gymnostomiella M. Fleisch., Gymnostomum Nees & Hornsch.,
Gyroweisia Schimp., Hennediella Paris, Hilpertia R. H. Zander,
Hymenostyliella E. B. Bartram, Hymenostylium Brid., Hyophila Brid.,

Hyophiladelphus (Müll. Hal.) R. H. Zander, Hypodontium Müll. Hal.,
Leptobarbula Schimp., Leptodontiella R. H. Zander & E. H. Hegew.,
Leptodontium (Müll. Hal.) Lindb., Luisierella Thér. & P. de la Varde,
Microbryum Schimp., Microcrossidium J. Guerra & M. J. Cano, Mironia
R. H. Zander, Molendoa Lindb., Neophoenix R. H. Zander & During,
Pachyneuropsis H. Mill., Phascopsis I. G. Stone, Plaubelia Brid., Pleurochaete
Lindb., Pottiopsis Blockeel & A. J. E. Sm., Pseudocrossidium R. S. Williams,
Pseudosymblepharis Broth., Pterygoneurum Jur., Quaesticula R. H. Zander,
Reimersia P. C. Chen, Rhexophyllum Herzog, Sagenotortula R. H. Zander,
Saitobryum R. H. Zander, Sarconeurum Bryhn, Scopelophila (Mitt.) Lindb.,
Splachnobryum Müll. Hal., Stegonia Venturi, Stonea R. H. Zander,
Streptocalypta Müll. Hal., Streptopogon Mitt., Streptotrichum Herzog,
Syntrichia Brid., Teniolophora W. D. Reese, Tetracoscinodon R. Br.,
Tetrapterum A. Jaeger, Timmiella (De Not.) Schimp., Tortella (Lindb.)
Limpr., Tortula Hedw. (Fig. 7), Trachycarpidium Broth., Trachyodontium
Steere, Trichostomum Bruch, Triquetrella Müll. Hal., Tuerckheimia Broth.,
Uleobryum Broth., Weisiopsis Broth., Weissia Hedw., Weissiodicranum W.
D. Reese, Willia Müll. Hal.
Ephemeraceae Schimp. Plants minute, developing from persistent
protonemata; capsules immersed, cleistocarpous or with poorly
differentiated operculum; columella disintegrating; Ephemerum
Schimp., Micromitrium Austin, Nanomitriopsis Cardot
Serpotortellaceae W. D. Reese & R. H. Zander. Plants robust,
cladocarpous, epiphytic; cauline central strand present; leaf margins
entire and unistratose; perichaetial leaves differentiated; peristome
well developed, reflexed when dry; Serpotortella Dixon
Calymperaceae Kindb. Plants epiphytic; stem lacking central strand;
leaves narrowly to broadly lanceolate; laminal cells papillose or
smooth; often with hyaline cancellinae on either side of costa at leaf
base; calyptrae persistent or not; peristome of 16 (rarely fused into 8)
segments, smooth, papillose or vertically striate; Arthrocormus Dozy &
Molk., Calymperes Sw., Exodictyon Cardot, Exostratum L. T. Ellis,
Leucophanes Brid., Mitthyridium H. Rob., Octoblepharum Hedw.,
Syrrhopodon Schwägr.
Cinclidotaceae K. Saito. Plants blackish and robust, generally
cladocarpous, riparian to aquatic; cauline central strand lacking;
leaves with thickened margins; capsules ribbed when dry; Cinclidotus
P. Beauv.
Bryobartramiaceae Sainsb. Plants very small, acrocarpous; calyptrae

remaining attached to vaginula, persisting as an epigonium; capsules
cleistocarpous; Bryobartramia Sainsb.
subclass: bryidae Peristome double, of alternating teeth and
segments; endostome ciliate; late stage division in the IPL asymmetric.
superorder: bryanae Plants acrocarpous or cladocarpous;
pseudoparaphyllia generally lacking; leaves erect to spreading,
lanceolate to ovate, mostly costate, costal anatomy mostly
heterogeneous; laminal cells generally short.
order: splachnales Laminal cells rhombic to elongate, typically smooth;
capsules erect with differentiated neck; peristome single or double; cilia
rudimentary or lacking.
Splachnaceae Grev. & Arn. Plants mostly coprophilous; laminal cells
thin-walled, rhomboidal; annulus not differentiated; capsules erect,
neck often differentiated into broad hypophysis; endostome fused to
exostome or lacking; Aplodon R. Br., Brachymitrion Taylor, Moseniella
Broth., Splachnum Hedw., Tayloria Hook., Tetraplodon Bruch &
Schimp., Voitia Hornsch.
Meesiaceae Schimp. Capsules more or less horizontal, with elongated
neck; annulus differentiated and revoluble; peristome double;
exostome typically shorter then endostome; Amblyodon P. Beauv.,
Leptobryum (Bruch & Schimp.) Wilson, Meesia Hedw., Neomeesia
Deguchi, Paludella Brid.
Catoscopiaceae Broth. Plants small, slender; leaves in three ranks;
laminal cells quadrate and smooth; capsules black, asymmetric,
horizontal; peristome double and reduced; Catoscopium Brid.
order: orthotrichales Plants medium-size, epiphytic or saxicolous;
cauline central strand lacking; laminal cells typically papillose; capsules
erect; peristome double or reduced; exostome recurved.
Orthotrichaceae Arn. Plants acrocarpous or cladocarpous; laminal
cells mostly isodiametric, thick-walled; calyptrae typically plicate
and hairy; capsules erect, rarely immersed, often ribbed; OPL thick
and teeth recurved when dry; cilia lacking; Bryomaltaea Goffinet,
Cardotiella Vitt, Ceuthotheca Lewinsky, Codonoblepharon Schwägr.,
Desmotheca Lindb., Florschuetziella Vitt, Groutiella Steere, Leiomitrium
Mitt., Leptodontiopsis Broth., Leratia Broth. & Paris, Macrocoma (Müll.
Hal.) Grout, Macromitrium Brid., Matteria Goffinet, Muelleriella Dusén,
Orthotrichum Hedw., Pentastichella Müll. Hal., Pleurorthotrichum Broth.,
Schlotheimia Brid., Sehnemobryum Lewinsky-Haapasaari & Hedenäs,
Stoneobryum D. H. Norris & H. Rob., Ulota D. Mohr, Zygodon Hook. &

Taylor
order: hedwigiales Plants medium to robust, plagiotropic, acrocarpous
or cladocarpous; laminal cells thick-walled, papillose or smooth; capsules
gymnostomous and immersed.
Hedwigiaceae Schimp. Protonemata globular; laminal cells
pluripapillose; calyptrae smooth, and naked; Braunia Bruch &
Schimp., Hedwigia P. Beauv., Hedwigidium Bruch & Schimp.,
Pseudobraunia (Lesq. & James) Broth.
Helicophyllaceae Broth. Leaves dimorphic, with lateral leaves
strongly inrolled when dry, dorsal and ventral leaves reduced and
appressed; laminal cells smooth; Helicophyllum Brid.
Rhacocarpaceae Kindb. Leaves bordered by narrow cells; laminal cells
roughened; alar cells inflated; Pararhacocarpus Frahm, Rhacocarpus
Lindb.
order: bryales Plants primarily terricolous; cauline central strand
present; laminal cells rhombic to elongate, smooth; annulus differentiated;
capsules pendent, neck differentiated; peristome double, typically well
developed and ciliate; exostome incurved.
Aulacomniaceae Schimp. Plants medium to robust; laminal cells
isodiametric, incrassate; capsules ribbed when dry; annulus
differentiated; peristome double, bryoid; Aulacomnium Schwägr.
Bartramiaceae Schwägr. Plants often robust; laminal cells
isodiametric, quadrate or rectangular, smooth or prorulose; annulus
typically undifferentiated; capsules subglose, erect or slightly curved,
typically ribbed; neck undifferentiated; Anacolia Schimp., Bartramia
Hedw., Breutelia (Bruch & Schimp.) Schimp., Conostomum Sw.,
Fleischerobryum Loeske, Flowersia D. G. Griffin & W. R. Buck, Leiomela
(Mitt.) Broth., Philonotis Brid., Plagiopus Brid., Quathlamba Magill
Orthodontiaceae (Broth.) Goffinet, comb. nov. (Bryaceae subfam.
Orthodontoideae Broth. in Engler, Die natürlichen
Pflanzenfamilien, ed. 2, 11: 347, 1924). Plants small to robust; laminal
cells elongate, lax; capsules often ribbed; annulus lacking;
endostomial membrane reduced or lacking; Orthodontium Wilson,
Orthodontopsis Ignatov & B. C. Tan
Bryaceae Schwägr. Plants erect, mostly unbranched, acrocarpous;
laminal cells mostly rhomboidal, smooth, thin-walled; costa single,
strong; capsules inclined to pendulous, smooth, with differentiated
neck; Acidodontium Schwägr., Anomobryum Schimp., Brachymenium
Schwägr., Bryum Hedw., Mielichhobryum J. P. Srivast., Mniobryoides

Hörmann, Perssonia Bizot, Plagiobryum Lindb., Pleurophascum Lindb.,
Rhodobryum (Schimp.) Limpr., Roellia Kindb., Rosulabryum J. R. Spence
Phyllodrepaniaceae Crosby. Plants small; leaves complanate, in four
rows; peristome single, of 16 segments; Mniomalia Müll. Hal.,
Phyllodrepanium Crosby.
Pseudoditrichaceae Steere & Z. Iwats. Plants very small; leaves ovate
lanceolate; laminal cells thick-walled; capsules erect; peristome
double; cilia lacking; Pseudoditrichum Steere & Z. Iwats.
Mniaceae Schwägr. Plants acro- or cladocarpous; leaves often bordered
and often toothed; laminal cells thin-walled, rhomboidal to
elongate; Cinclidium Sw., Cyrtomnium Holmen, Epipterygium Lindb.,
Leucolepis Lindb., Mielichhoferia Nees & Hornsch., Mnium Hedw.,
Orthomnion Wilson, Plagiomnium T. J. Kop., Pohlia Hedw., Pseudobryum
(Kindb.) T. J. Kop., Pseudopohlia R. S. Williams, Rhizomnium (Broth.)
T. J. Kop., Schizymenium Harv., Synthetodontium Cardot, Trachycystis T. J.
Kop.
Leptostomataceae Schwägr. Plants forming dense mats; stems heavily
tomentose; leaf margins entire, unbordered; annulus poorly
differentiated to lacking; stomata cryptoporous; peristome strongly
reduced; Leptostomum R. Br.
superorder: rhizogonianae Plants erect, generally from
rhizomatous stem, acro-, clado-, or pleurocarpous; leaves oblong to
lanceolate, costate, costal anatomy heterogeneous; laminal cells
generally short and thick-walled.
order: rhizogoniales
Hypnodendraceae Broth. Plants robust, rhizomatous and stipitate;
secondary stems erect, frondose to dendroid; laminal marginal cells
differentiated or not, unistratose, often toothed; capsules often
ribbed when dry; annulus differentiated; peristome well developed,
ciliate; Hypnodendron (Müll. Hal.) Mitt.
Rhizogoniaceae Broth. Plants medium to large; cauline central strand
present; marginal laminal cells often differentiated, bi- or
multistratose, often toothed; costa typically toothed above; annulus
differentiated; capsules generally smooth; peristome typically well
developed, ciliate, or reduced to endostome or exostome;
Cryptopodium Brid., Goniobryum Lindb., Hymenodon Hook.f. & Wilson,
Hymenodontopsis Herzog, Leptotheca Schwägr., Mesochaete Lindb.,
Pyrrhobryum Mitt., Rhizogonium Brid.
Calomniaceae Kindb. Plants small, arising from persistent protonema;

leaves tristichous and dimorphic, with ventral ones much smaller;
perichaetial leaves highly differentiated; capsules gymnostomous;
Calomnion Hook. f. & Wilson
Mitteniaceae Broth. Plants small, with luminescent protonemata;
cauline central strand lacking; leaves complanate, decurrent;
perichaetia polysetous; peristome double, outer row homologous to
bryoid endostome; Mittenia Lindb.
Cyrtopodaceae M. Fleisch. Plants robust and large, with rhizomatous
stem; secondary stems erect, frondose to dendroid; lamina bistratose
above; margin unistratose, toothed or entire; costa toothed above;
capsules smooth; peristome double or reduced, eciliate; Bescherellia
Duby, Cyrtopodendron M. Fleisch., Cyrtopus (Brid.) Hook. f.
Spiridentaceae Kindb. Plants robust and large, with rhizomatous
stem; secondary stems erect, sparsely branched; lamina bi- or
pluristratose; margin typically toothed; costa toothed above;
capsules smooth, immersed or exserted; peristome double, teeth and
segments long, eciliate; Franciella Thér., Spiridens Nees
Pterobryellaceae (Broth.) W. R. Buck & Vitt. Plants robust and large,
stipitate from rhizomatous stem, frondose to dendroid; cauline
central strand lacking; capsules short oval; annulus differentiated;
peristome double, with long teeth and segments but reduced cilia;
Pterobryella (Müll. Hal.) A. Jaeger
Racopilaceae Kindb. Stems plagiotropic; leaves dimorphic with dorsal
ones reduced; costa excurrent; capsules long–exserted; peristome
double, well developed; Powellia Mitt., Racopilum P. Beauv.
superorder: hypnanae Plants pleurocarpous, typically freely
branching; pseudoparaphyllia usually present; leaves mostly ovate,
costate or not, costal anatomy homogeneous; laminal cells generally
elongate.
order: hookeriales Alar cells mostly not differentiated; opercula mostly
rostrate; exostome teeth often furrowed.
suborder: Hookeriineae Plants mostly medium-sized to slender, never
(?) phyllodioicous; stems monopodially branched; leaves seldom plicate;
laminal cells often large and lax, thin-walled; costae often strong and
double, sometimes single or short and double; capsules mostly smooth;
exothecial cells often collenchymatous; opercula often rostrate; exostome
teeth often furrowed; endostomial segments with baffle-like cross-walls;
calyptrae often mitrate.
Hookeriaceae Schimp. Stems with central strand; pseudoparaphyllia

filamentous or absent; laminal cells large and lax; costa usually short
and double; calyptrae multistratose at middle, naked; Achrophyllum Vitt
& Crosby, Cyathophorella (Broth.) M. Fleisch., Cyathophorum P. Beauv.,
Dendrocyathophorum Dixon, Hookeria J. E. Sm., Schimperobryum Margad.
Leucomiaceae Broth. Stems lacking central strand; pseudoparaphyllia
absent; laminal cells linear, lax; costa lacking; calyptrae cucullate;
Leucomium Mitt., Philophyllum Müll. Hal., Rhynchostegiopsis Müll. Hal.
Daltoniaceae Schimp. Stems lacking central strand; pseudoparaphyllia
absent; laminal cells oval to long–hexagonal, differentiated at leaf
margins; costa single; calyptrae unistratose at middle, fringed at
base, usually naked; Calyptrochaeta Desv., Crosbya Vitt, Daltonia Hook.
& Taylor, Distichophyllidium M. Fleisch., Distichophyllum Dozy &
Molk., Ephemeropsis K. I. Goebel, Leskeodon Broth., Leskeodontopsis
Zanten, Metadistichophyllum Nog. & Z. Iwats.
Adelotheciaceae W. R. Buck. Stems lacking central strand;
pseudoparaphyllia filamentous; laminal cells isodiametric, not
differentiated at leaf margins; costa single; calyptrae unistatose at
middle, not fringed at base, densely hairy; Adelothecium Mitt.,
Bryobrothera Thér.
Pilotrichaceae Kindb. Stems lacking central strand; pseudoparaphyllia
none or foliose; laminal cells various; costa strong and double, or
short and double; calyptrae unistratose at middle, usually hairy;
Actinodontium Schwägr., Amblytropis (Mitt.) Broth., Beeveria Fife,
Brymela Crosby & B. H. Allen (Fig. 3), Callicostella (Müll. Hal.) Mitt.,
Callicostellopsis Broth., Crossomitrium Müll. Hal., Cyclodictyon Mitt.,
Diploneuron E. B. Bartram, Helicoblepharum (Mitt.) Broth., Hemiragis
(Brid.) Besch., Hookeriopsis (Besch.) A. Jaeger, Hypnella (Müll. Hal.) A.
Jaeger, Lepidopilidium (Müll. Hal.) Broth., Lepidopilum (Brid.) Brid.,
Pilotrichidium Besch., Pilotrichum P. Beauv., Sauloma (Hook. f. & Wilson)
Mitt., Stenodesmus (Mitt.) A. Jaeger, Stenodictyon (Mitt.) A. Jaeger,
Tetrastichium (Mitt.) Cardot, Thamniopsis (Mitt.) M. Fleisch.,
Trachyxiphium W. R. Buck, Vesiculariopsis Broth.
Hypopterygiaceae Mitt. Plants dendroid; amphigastria differentiated;
leaves often limbate; costa single; laminal cells short, mostly smooth;
alar cells not differentiated; exostome teeth not furrowed;
endostomial segments lacking baffle-like cross-walls;
Canalohypopterygium W. Frey & Schaepe, Catharomnion Hook. f. &
Wilson, Hypopterygium Brid., Lopidium Hook. f. & Wilson
suborder: Ptychomniieae Plants usually robust and turgid, often

phyllodioicous; stems sympodially branched, usually lacking a central
strand; leaves usually plicate, often strongly toothed; costae short and
double; laminal cells elongate, often thick-walled and porose; alar cells
often colored; capsules mostly ribbed; endostomial segments lacking
baffle-like cross-walls; calyptrae often cucullate.
Ptychomniaceae M. Fleisch. Alar cells mostly little differentiated,
except for color; capsules long–exserted, strongly eight-ribbed;
calyptrae cucullate; spores isosporous; Cladomnion Hook. f. & Wilson,
Cladomniopsis M. Fleisch., Glyphothecium Hampe, Hampeella Müll.
Hal., Ptychomnium (Hook.f. & Wilson) Mitt., Tetraphidopsis Broth. &
Dixon
Garovagliaceae (M. Fleisch.) W. R. Buck & Vitt. Alar cells typically well
differentiated in shape and color; capsules often immersed, smooth
or somewhat sulcate; calyptrae mitrate or cucullate; spores
anisosporous; Endotrichellopsis During, Euptychium Schimp.,
Garovaglia Endl.
order: hypnales Stems monopodially or sympodially branched; alar cells
often differentiated; opercula various, mostly not rostrate; exostome seldom
furrowed; calyptrae mostly cucullate, naked.
Rutenbergiaceae (Broth.) M. Fleisch. Stems sympodially branched,
lacking a central strand; secondary stems little branched; costa
single; laminal cells prorulose; alar cells well differentiated; capsules
immersed; calyptrae mitrate, hairy; Neorutenbergia Bizot & Pócs,
Pseudocryphaea Broth., Rutenbergia Besch.
Trachylomataceae (M. Fleisch.) W. R. Buck & Vitt. Stems sympodially
branched; secondary stems stipitate frondose, complanate–foliate;
alar cells weakly differentiated; asexual propagula of stem-borne,
filamentous gemmae; exostome teeth pale, densely papillose;
Braithwaitea Lindb., Trachyloma Brid.
Amblystegiaceae G. Roth. Plants typically growing in moist areas;
stems monopodially branched; costa often variable; laminal cells
mostly short; alar cells not or weakly differentiated; setae relatively
long in comparison to size of plants; capsules strongly curved and
asymmetric; exostome teeth yellow-brown, cross-striolate;
Amblystegium Schimp., Donrichardsia H. A. Crum & L. E. Anderson,
Gradsteinia Ochyra, Hygroamblystegium Loeske, Hypnobartlettia Ochyra,
Koponenia Ochyra, Leptodictyum (Schimp.) Warnst., Limbella (Müll.
Hal.) Müll. Hal., Ochyraea Váňa, Orthotheciella (Müll. Hal.) Ochyra,
Platylomella A. L. Andrews, Sciaromiella Ochyra, Sciaromiopsis Broth.,

Sinocalliergon Sakurai, Vittia Ochyra
Cratoneuraceae Mönk. Stems monopodially branched, often wiry,
often densely tomentose; paraphyllia present, foliose; stem and
branch leaves differentiated; costa single; laminal cells relatively
short, smooth or prorulose; alar cells well differentiated; exostome
teeth yellow-brown, cross-striolate; Cratoneuron (Sull.) Spruce,
Sasaokaea Broth.
Helodiaceae (M. Fleisch.) Ochyra. Stems monopodially branched;
paraphyllia present, filamentous to narrowly foliose, the cells
elongate, not papillose; costa single; laminal cells mostly prorulose;
alar cells often well differentiated; exostome teeth cross-striolate;
Actinothuidium (Besch.) Broth., Bryochenea C. H. Gao & K. C. Chang,
Helodium Warnst. (Fig. 2), Palustriella Ochyra
Hylocomiaceae (Broth.) M. Fleisch. Plants mostly robust; stems
monopodially or sympodially branched; paraphyllia often present;
leaves often strongly toothed; costae often strong and double;
laminal cells elongate, smooth or prorulose; alar cells weakly
differentiated; setae very elongate; capsules typically curved and
asymmetric; exostome teeth yellow- to red-brown, often with
reticulate pattern; Hylocomiastrum Broth., Hylocomium Bruch &
Schimp., Leptocladiella M. Fleisch., Leptohymenium Schwägr.,
Loeskeobryum Broth., Macrothamnium M. Fleisch., Meteoriella S.
Okamura, Neodolichomitra Nog., Orontobryum M. Fleisch., Pleurozium
Mitt., Rhytidiadelphus (Limpr.) Warnst., Rhytidiopsis Broth.,
Schofieldiella W. R. Buck
Rhytidiaceae Broth. Plants robust; stems monopodially branched;
paraphyllia none; leaves plicate, rugose; costa single; laminal cells
linear, strongly porose, prorulose; alar cells well differentiated;
exostome teeth yellow-brown, cross-striolate; Rhytidium (Sull.)
Kindb.
Leskeaceae Schimp. Plants terrestrial or epiphytic; stems
monopodially branched, often terete–foliate; paraphyllia
nonpapillose; leaves mostly short–acuminate; costa mostly single;
laminal cells short, usually unipapillose; alar cells weakly
differentiated; capsules curved and asymmetric when plants
terrestrial but in epiphytes often erect; exostome striate in terrestrial
taxa but in epiphytes often pale, weakly ornamented; endostome
often reduced; Claopodium (Lesq. & James) Renauld & Cardot,
Dolichomitriopsis S. Okamura, Fabronidium Müll. Hal., Haplocladium

(Müll. Hal.) Müll. Hal., Hylocomiopsis Cardot, Leptocladium Broth.,
Leptopterigynandrum Müll. Hal., Lescuraea Bruch & Schimp., Leskea
Hedw., Leskeadelphus Herzog, Leskeella (Limpr.) Loeske, Lindbergia
Kindb., Mamillariella Laz., Miyabea Broth., Okamuraea Broth.,
Orthoamblystegium Dixon & Sakurai, Pseudoleskea Bruch & Schimp.,
Pseudoleskeella Kindb., Pseudoleskeopsis Broth., Ptychodium Schimp.,
Rigodiadelphus Dixon, Schwetschkea Müll. Hal.
Regmatodontaceae Broth. Plants epiphytic; stems monopodially
branched; paraphyllia absent; costa single; laminal cells short, smooth;
alar cells not or weakly, differentiated; capsules erect; exostome teeth
much shorter than endostome segments; Regmatodon Brid.
Pterigynandraceae Schimp. Plants terrestrial or epiphytic, mostly
relatively small; stems monopodially branched, mostly
terete–foliate; paraphyllia absent; costa short and double; laminal
cells short, prorulose; alar cells weakly differentiated; gemmae stem-
borne; setae smooth; capsules often erect; peristome often reduced;
Habrodon Schimp., Heterocladium Bruch & Schimp., Iwatsukiella W. R.
Buck & H. A. Crum, Myurella Bruch & Schimp., Pterigynandrum Hedw.,
Trachyphyllum A. Gepp
Rigodiaceae H. A. Crum. Plants terrestrial or weakly epiphytic, more
or less stipitate; stems monopodially branched; paraphyllia absent;
stipe, stem and branch leaves differentiated; costa single; laminal
cells short, smooth; alar cells not or weakly differentiated; setae
smooth; capsules curved and asymmetric; exostome teeth densely
cross-striolate; Rigodium Schwägr.
Thuidiaceae Schimp. Plants terrestrial; stems monopodially branched;
paraphyllia present, the cells papillose; stem and branch leaves
differentiated; costa single; laminal cells short, papillose; alar cells
not or weakly differentiated; setae often roughened; capsules
typically curved and asymmetric; exostome teeth densely cross-
striolate; calyptrae naked or sparsely hairy.; Abietinella Müll. Hal.,
Boulaya Cardot, Cyrto-hypnum (Hampe) Hampe & Lorentz, Fauriella
Besch., Pelekium Mitt., Rauiella Reimers, Thuidiopsis (Broth.) M.
Fleisch., Thuidium Bruch & Schimp.
Campyliaceae (Kanda) W. R. Buck, comb. nov. (Amblystegiaceae
subfam. Campylioideae Kanda, J. Sci. Hiroshima Univ., Ser. B, Div. 2,
Bot. 15: 250. 1975 [1976]). Plants typically growing in moist areas,
relatively robust; stems monopodially branched; costa mostly single;
laminal cells elongate; alar cells often differentiated; setae not

especially long; capsules asymmetric; exostome teeth yellow-brown;
Anacamptodon Brid., Callialaria Ochyra, Calliergon (Sull.) Kindb.,
Campyliadelphus (Kindb.) R. S. Chopra, Campylium (Sull.) Mitt.,
Conardia H. Rob., Cratoneuropsis (Broth.) M. Fleisch., Drepanocladus
(Müll. Hal.) G. Roth, Hamatocaulis Hedenäs, Hygrohypnum Lindb.,
Loeskypnum H. K. G. Paul, Pictus C. C. Towns., Pseudocalliergon (Limpr.)
Loeske, Sanionia Loeske, Scorpidium (Schimp.) Limpr., Straminergon
Hedenäs, Tomentypnum Loeske, Warnstorfia Loeske
Brachytheciaceae G. Roth. Plants mostly growing in mesic woodlands,
terrestrial; stems monopodially branched; leaves often plicate; costa
single, often projecting as a small spine; laminal cells elongate; alar
cells mostly weakly differentiated; setae sometimes roughened;
capsules often relatively short, curved, asymetric; opercula conic to
rostrate; exostome teeth mostly red-brown; calyptrae mostly naked;
Aerobryum Dozy & Molk., Aerolindigia M. Menzel, Bestia Broth.,
Brachythecium Schimp., Bryhnia Kaurin, Bryoandersonia H. Rob.,
Bryostreimannia Ochyra, Cirriphyllum Grout, Clasmatodon Hook. f. &
Wilson, Cratoneurella H. Rob., Eriodon Mont., Eurhynchiella M. Fleisch.,
Eurhynchium Bruch & Schimp., Flabellidium Herzog, Homalotheciella
(Cardot) Broth., Homalothecium Schimp., Isothecium Brid., Juratzkaeella
W. R. Buck, Lindigia Hampe, Mandoniella Herzog, Meteoridium (Müll.
Hal.) Manuel, Myuroclada Besch., Nobregaea Hedenäs, Palamocladium
Müll. Hal., Plasteurhynchium Broth., Platyhypnidium M. Fleisch.,
Pseudopleuropus Takaki, Pseudoscleropodium (Limpr.) M. Fleisch. (Fig. 8),
Puiggariopsis M. Menzel, Rhynchostegiella (Schimp.) Limpr.,
Rhynchostegium Bruch & Schimp., Rozea Besch., Schimperella Thér.,
Scleropodiopsis Ignatov, Scleropodium Bruch & Schimp., Scorpiurium
Schimp., Squamidium (Müll. Hal.) Broth., Steerecleus H. Rob.,
Stenocarpidiopsis M. Fleisch., Trachybryum (Broth.) W. B. Schofield,
Zelometeorium Manuel
Stereophyllaceae (M. Fleisch.) W. R. Buck & Ireland. Plants terrestrial
or epiphytic; stems monopodially branched; costa typically single;
laminal cells elongate, mostly smooth, rarely unipapillose; alar cells
differentiated, collenchymatous, extending across base of costa; setae
smooth; capsules inclined to erect; exostome teeth cross-striolate to
papillose; Catagoniopsis Broth., Entodontopsis Broth., Eulacophyllum W.
R. Buck & Ireland, Juratzkaea Lorentz, Pilosium (Müll. Hal.) M. Fleisch.,
Sciuroleskea Broth., Stenocarpidium Müll. Hal., Stereophyllum Mitt.
Myriniaceae Schimp. Plants often epiphytic, small; stems

monopodially branched; costa single, often slender; laminal cells
elongate, smooth; alar cells weakly differentiated; capsules often
erect; peristomes mostly variously reduced; calyptrae rarely hairy;
Austinia Müll. Hal., Helicodontium Schwägr., Macgregorella E. B.
Bartram, Merrilliobryum Broth., Myrinia Schimp., Nematocladia W. R.
Buck
Fabroniaceae Schimp. Plants epiphytic, often small; stems
monopodially branched, sometimes fragile; leaves mostly
acuminate; costa single, slender; laminal cells short, smooth; alar
cells mostly weakly differentiated; capsules typically erect; peristome
often reduced; exostome teeth often paired; Dimerodontium Mitt.,
Fabronia Raddi, Ischyrodon Müll. Hal., Levierella Müll. Hal.,
Rhizofabronia (Broth.) M. Fleisch.
Meteoriaceae Kindb. Plants epiphytic, often pendent; stems
monopodially branched, often very elongate; costa short and double
or single; laminal cells mostly elongate, sometimes short, often
variously papillose; alar cells not or weakly differentiated; setae often
short, roughened; capsules often immersed, erect, symmetric;
exostome teeth cross-striolate to papillose; endostome often
reduced; calyptrae mitrate or cucullate, often hairy; Aerobryidium M.
Fleisch., Aerobryopsis M. Fleisch., Ancistrodes Hampe, Barbella M.
Fleisch., Barbellopsis Broth., Chrysocladium M. Fleisch., Cryphaeophilum
M. Fleisch., Cryptopapillaria M. Menzel, Diaphanodon Renauld &
Cardot, Duthiella Renauld, Floribundaria M. Fleisch., Lepyrodontopsis
Broth., Meteoriopsis Broth., Meteorium (Brid.) Dozy & Molk.,
Neodicladiella W. R. Buck, Neonoguchia S. H. Lin, Pseudospiridentopsis
(Broth.) M. Fleisch., Pseudotrachypus P. de la Varde & Thér., Sinskea W.
R. Buck, Toloxis W. R. Buck, Trachycladiella (M. Fleisch.) M. Menzel &
W. Schultze-Motel, Trachypodopsis M. Fleisch., Trachypus Reinw. &
Hornsch.
Plagiotheciaceae (Broth.) M. Fleisch. Plants terrestrial; stems
monopodially branched, mostly complanate–foliate; leaves
decurrent; costa short and double or absent; laminal cells elongate,
often strongly chlorophyllose; alar cells differentiated into the
decurrencies; setae smooth; capsules often curved and asymmetric;
peristome teeth mostly pale yellow; exostome typically cross-
striolate below; endostome well developed; Plagiothecium Bruch &
Schimp.
Fontinalaceae Schimp. Plants aquatic; stems sympodially branched;

costa single or short and double (and then the leaves concave to
carinate); capsules immersed or short–exserted; endostome forming
a trellis; calyptrae mitrate or cucullate; Brachelyma Cardot, Dichelyma
Myrin, Fontinalis Hedw.
Entodontaceae Kindb. Plants often epiphytic; stems monopodially
branched; costa short and double or absent; laminal cells linear,
smooth; alar cells subquadrate, numerous; capsules erect and
symmetric, long–exserted; columella often exserted; peristome
inserted below mouth of capsule; endostome mostly strongly
reduced; Entodon Müll. Hal., Erythrodontium Hampe, Mesonodon
Hampe, Pylaisiobryum Broth.
Climaciaceae Kindb. Plants dendroid; stems sympodially branched,
with paraphyllia on stipe; leaves not decurrent; costa single; laminal
cells relatively short, smooth; capsules erect and symmetric;
Climacium F. Weber & D. Mohr
Pleuroziopsidaceae Ireland. Plants dendroid; stems sympodially
branched, with longitudinal lamellae on stipe; leaves decurrent;
costa single; laminal cells relatively short, smooth; capsules inclined
and asymmetric; Pleuroziopsis E. Britton
Hypnaceae Schimp. Stems monopodially branched; pseudoparaphyllia
foliose or rarely filamentous; paraphyllia none; leaves often falcate or
homomallous; costa short and double (or absent); laminal cells
mostly linear; capsules mostly inclined and asymmetric; exothecial
cells usually not collenchymatous; opercula apiculate to
short–rostrate; exostome teeth mostly cross-striolate; calyptrae
mostly naked; Andoa Ochyra, Bardunovia Ignatov & Ochyra, Breidleria
Loeske, Bryocrumia L. E. Anderson, Bryosedgwickia Cardot & Dixon,
Callicladium H. A. Crum, Calliergonella Loeske, Campylophyllum
(Schimp.) M. Fleisch., Caribaeohypnum Ando & Higuchi, Chryso-
hypnum (Hampe) Hampe, Crepidophyllum Herzog, Ctenidiadelphus M.
Fleisch., Ctenidium (Schimp.) Mitt., Cyathothecium Dixon,
Ectropotheciella M. Fleisch., Ectropotheciopsis (Broth.) M. Fleisch.,
Ectropothecium Mitt., Elharveya H. A. Crum, Elmeriobryum Broth.,
Entodontella M. Fleisch., Eurohypnum Ando, Fallaciella H. A. Crum,
Giraldiella Müll. Hal., Gollania Broth., Herzogiella Broth., Homomallium
(Schimp.) Loeske, Hondaella Dixon & Sakurai, Horridohypnum W. R.
Buck, Hyocomium Bruch & Schimp., Hypnum Hedw., Irelandia W. R.
Buck, Isopterygiopsis Z. Iwats., Isopterygium Mitt., Leiodontium Broth.,
Macrothamniella M. Fleisch., Mahua W. R. Buck, Microctenidium M.

Fleisch., Mittenothamnium Henn., Nanothecium Dixon & P. de la Varde,
Orthothecium Bruch & Schimp., Phyllodon Bruch & Schimp.,
Plagiotheciopsis Broth., Platydictya Berk., Platygyriella Cardot,
Platygyrium Bruch & Schimp., Podperaea Z. Iwats. & Glime,
Pseudotaxiphyllum Z. Iwats., Ptilium De Not., Pylaisiella Grout,
Rhacopilopsis Renauld & Cardot, Rhizohypnella M. Fleisch.,
Sclerohypnum Dixon, Serpoleskea (Limpr.) Loeske, Stenotheciopsis Broth.,
Stereodontopsis R. S. Williams, Syringothecium Mitt., Taxiphyllopsis
Higuchi & Deguchi, Taxiphyllum M. Fleisch., Trachythecium
M. Fleisch., Tripterocladium (Müll. Hal.) A. Jaeger, Vesicularia (Müll.
Hal.) Müll. Hal., Wijkiella Bizot & Lewinsky
Catagoniaceae W. R. Buck & Ireland. Stems monopodially branched;
pseudoparaphyllia filamentous; leaves conduplicate; costa short and
double or absent; laminal cells linear, smooth; alar cells not
differentiated; exostome teeth cross-striolate; Catagonium Broth.
Symphyodontaceae M. Fleisch. Stems monopodially branched;
laminal cells mostly prorulose; alar cells not or weakly differentiated;
setae mostly roughened; capsules symmetric, typically spinose;
exostome teeth papillose to cross-striolate; calyptrae cucullate or
mitrate; Chaetomitriopsis M. Fleisch., Chaetomitrium Dozy & Molk.,
Dimorphocladon Dixon, Symphyodon Mont.
Sematophyllaceae Broth. Stems monopodially branched; leaves often
golden green; costa short and double or none; laminal cells mostly
linear, smooth or papillose; alar cells well differentiated; exothecial
cells mostly collenchymatous; opercula mostly obliquely rostrate;
exostome teeth often furrowed, cross-striolate; Acanthorrhynchium M.
Fleisch., Acritodon H. Rob., Acroporium Mitt., Allioniellopsis Ochyra,
Aptychella (Broth.) Herzog, Aptychopsis (Broth.) M. Fleisch., Brotherella
M. Fleisch., Chinostomum Müll. Hal., Clastobryella M. Fleisch.,
Clastobryophilum M. Fleisch., Clastobryopsis M. Fleisch., Clastobryum
Dozy & Molk., Colobodontium Herzog, Donnellia Austin, Foreauella
Dixon & P. de la Varde, Gammiella Broth., Hageniella Broth.,
Heterophyllium (Schimp.) Kindb., Isocladiella Dixon, Leptoischyrodon
Dixon, Macrohymenium Müll. Hal., Mastopoma Cardot, Meiotheciella B.
C. Tan, W. B. Schofield & H. P. Ramsay, Meiothecium Mitt., Papillidiopsis
(Broth.) W. R. Buck & B. C. Tan, Paranapiacabaea W. R. Buck & Vital,
Potamium Mitt., Pseudohypnella (M. Fleisch.) Broth., Pterogonidium
Broth., Pterogoniopsis Müll. Hal., Pylaisiadelpha Cardot, Pylaisiopsis
(Broth.) Broth., Radulina W. R. Buck & B. C. Tan, Rhaphidostichum M.

Fleisch., Schraderella Müll. Hal., Schroeterella Herzog, Sematophyllum
Mitt., Struckia Müll. Hal., Taxitheliella Dixon, Taxithelium Mitt.,
Timotimius W. R. Buck, Trichosteleum Mitt., Trismegistia (Müll. Hal.)
Müll. Hal., Trolliella Herzog, Warburgiella Müll. Hal., Wijkia H. A.
Crum
Hydropogonaceae W. H. Welch. Plants aquatic; stems monopodially
branched; costa short and double or none; laminal cells linear,
smooth; alar cells weakly differentiated; capsules cladocarpous,
immersed; endostome none; calyptrae mitrate; Hydropogon Brid.,
Hydropogonella Cardot
Myuriaceae M. Fleisch. Stems sympodially branched; secondary stems
little or not branched; leaves mostly long–acuminate; costa short and
double or none; laminal cells linear, smooth; alar cells well
differentiated, mostly colored; capsules long–exserted, erect;
exostome teeth reduced, smooth, often perforate; endostome
rudimentary; Eumyurium Nog., Myurium Schimp., Oedicladium Mitt.,
Palisadula Toyama, Piloecium (Müll. Hal.) Broth.
Cryphaeaceae Schimp. Stems sympodially branched; secondary stems
little or not branched; costa single; laminal cells short, smooth or
sometimes prorulose; alar cells numerous; capsules immersed or
seldom emergent; exostome teeth pale, papillose; endostome
rudimentary to absent; calyptrae mitrate; Cryphaea D. Mohr,
Cryphidium (Mitt.) A. Jaeger, Cyptodon (Broth.) M. Fleisch.,
Cyptodontopsis Dixon, Dendroalsia E. Britton, Dendrocryphaea Broth.,
Dendropogonella E. Britton, Pilotrichopsis Besch., Schoenobryum Dozy &
Molk., Sphaerotheciella M. Fleisch.
Prionodontaceae Broth. Plants epiphytic; stems sympodially
branched; axillary hairs as in Breutelia (Bartramiaceae); leaves usually
plicate and with strongly toothed margins; costa single; laminal cells
short, papillose; alar cells differentiated in large areas; capsules
immersed to emergent; annulus revoluble; exostome teeth papillose;
endostome segments united into a reticulum; Prionodon Müll. Hal.
Leucodontaceae Schimp. Plants mostly epiphytic; stems sympodially
branched; secondary stems often not or little branched, mostly curled
when dry; leaves rapidly spreading when moist, mostly plicate; costa
short and double or none; laminal cells oval to linear, mostly smooth,
rarely prorulose; alar cells numerous; capsules usually exserted;
annulus not differentiated; exostome teeth pale, papillose;
endostome mostly rudimentary; spores often large and

anisosporous; Antitrichia Brid., Dozya Sande Lac., Eoleucodon H. A.
Mill. & H. Whittier, Felipponea Broth., Leucodon Schwägr., Pterogonium
Sw., Scabridens E. B. Bartram
Pterobryaceae Kindb. Plants mostly epiphytic; stems sympodially
branched; secondary stems often well branched, and thus stipitate;
stem and branch leaves often differentiated, branch leaves sometimes
five-seriate; costa mostly single, sometimes short and double or
absent; laminal cells mostly linear, mostly smooth, sometimes
prorulose; alar cells usually differentiated, often thick-walled and
colored; capsules mostly immersed; exostome teeth pale, often
smooth; endostome most rudimentary; calyptrae cucullate or
mitrate, often hairy; Calyptothecium Mitt., Cryptogonium (Müll. Hal.)
Hampe, Henicodium (Müll. Hal.) Kindb., Hildebrandtiella Müll. Hal.,
Horikawaea Nog., Jaegerina Müll. Hal., Micralsopsis W. R. Buck,
Muellerobryum M. Fleisch., Neolindbergia M. Fleisch., Orthorrhynchidium
Renauld & Cardot, Orthostichidium Dusén, Orthostichopsis Broth.,
Osterwaldiella Broth., Penzigiella M. Fleisch., Pireella Cardot,
Pseudopterobryum Broth., Pterobryidium Broth. & Watts, Pterobryon
Hornsch., Pterobryopsis M. Fleisch., Pulchrinodus B. H. Allen, Renauldia
Müll. Hal., Rhabdodontium Broth., Spiridentopsis Broth., Symphysodon
Dozy & Molk., Symphysodontella M. Fleisch.
Phyllogoniaceae Kindb. Plants epiphytic; stems sympodially
branched; secondary stems irregularly branched, strongly
complanate–foliate; leaves conduplicate, cucullate, auriculate; costa
short and double or absent; laminal cells linear, smooth; alar cells
differentiated in small groups; capsules immersed or shortly
exserted; exostome teeth pale, not or scarcely ornamented;
endostome rudimentary or absent; calyptrae cucullate or mitrate,
naked or hairy; Phyllogonium Brid.
Orthorrhynchiaceae S. H. Lin. Plants terrestrial; stems monopodially
branched; leaves conduplicate, cucullate; costa short and double or
absent; laminal cells linear, smooth; alar cells undifferentiated;
capsules short–exserted, erect; exostome teeth pale, unornamented;
endostome none; calyptrae mitrate, hairy; Orthorrhynchium
Reichardt
Lepyrodontaceae Broth. Plants terrestrial or epiphytic; stems
sympodially branched; secondary stems not or little branched; leaves
sometimes plicate; costa single and weak or short and double to
absent; laminal cells linear, smooth, thick-walled and porose; alar

cells few or scarcely differentiated; capsules long–exserted;
peristome usually only endostomial; Dichelodontium Broth., Lepyrodon
Hampe
Neckeraceae Schimp. Plants terrestrial or epiphytic; stems mostly
sympodially branched, sometimes monopodial; stipes sometimes
differentiated and plants then frondose; leaves mostly complanately
arranged; costa typically single, sometimes short and double;
laminal cells fusiform to linear, rarely shorter, mostly smooth, rarely
prorulose or papillose; alar cells mostly few or weakly differentiated;
capsules immersed (mostly in epiphytes) to long–exserted (mostly in
terrestrial taxa); exostome teeth often pale, usually cross-striolate at
least at extreme base, papillose above; endostome often reduced;
calyptrae mostly cucullate; Baldwiniella M. Fleisch., Bissetia Broth.,
Bryolawtonia D. H. Norris & Enroth, Caduciella Enroth, Crassiphyllum
Ochyra, Cryptoleptodon Renauld & Cardot, Curvicladium Enroth,
Dixonia Horik. & Ando, Dolichomitra Broth., Handeliobryum Broth.,
Himantocladium (Mitt.) M. Fleisch., Homalia (Brid.) Bruch & Schimp.,
Homaliadelphus Dixon & P. de la Varde, Homaliodendron M. Fleisch.,
Hydrocryphaea Dixon, Isodrepanium (Mitt.) E. Britton, Metaneckera
Steere, Neckera Hedw., Neckeropsis Reichardt, Neomacounia Ireland,
Noguchiodendron Ninh & Pócs, Pendulothecium Enroth & S. He,
Pinnatella M. Fleisch., Porothamnium M. Fleisch., Porotrichodendron M.
Fleisch., Porotrichopsis Broth. & Herzog, Porotrichum (Brid.) Hampe,
Thamnobryum Nieuwl., Touwia Ochyra
Echinodiaceae Broth. Plants epipetric or less often on soil or bases of
trees; stems sympodially branched, wiry; secondary stems irregularly
branched; leaves mostly subulate; costa single, mostly excurrent;
laminal cells short, smooth; alar cells weakly differentiated; capsules
long–exserted, inclined to horizontal; exostome teeth reddish, cross-
striolate; endostome well developed; Echinodium Jur.
Leptodontaceae Schimp. Plants mostly epiphytic; stems sympodially
branched, often curled when dry; secondary stems irregularly
branched to bipinnate; costa typically single; laminal cells
isodiametric to long–hexagonal, smooth, unipapillose or prorulose;
alar cells numerous; capsules immersed to short–exserted; exostome
teeth pale, unornamented to spiculose; endostome rudimentary;
calyptrae hairy; Alsia Sull., Forsstroemia Lindb., Leptodon D. Mohr,
Taiwanobryum Nog.
Lembophyllaceae Broth. Plants often turgid; stems monopodially

branched; leaves mostly strongly concave; costa mostly short and
double (rarely single); laminal cells elongate, smooth; alar cells often
somewhat differentiated; capsules mostly erect and immersed to
short–exserted; endostome mostly reduced; calyptrae rarely mitrate,
naked or hairy; Acrocladium Mitt., Camptochaete Reichardt, Fifea H. A.
Crum, Lembophyllum Lindb., Neobarbella Nog., Orthostichella Müll.
Hal., Pilotrichella (Müll. Hal.) Besch., Weymouthia Broth.
Anomodontaceae Kindb. Plants mostly epiphytic; stems sympodially
or monopodially branched, secondary stems and/or branches often
curled when dry, not complanate–foliate; paraphyllia none; leaves
often acute to obtuse; costa single or short and double; laminal cells
mostly short, papillose or prorulose; alar cells mostly poorly
differentiated; capsules exserted, erect; exostome teeth pale to white,
cross-striolate sometimes with overlying papillae to papillose;
endostome often reduced; Anomodon Hook. & Taylor, Bryonorrisia L. R.
Stark & W. R. Buck, Chileobryon Enroth, Curviramea H. A. Crum,
Haplohymenium Dozy & Molk., Herpetineuron (Müll. Hal.) Cardot,
Schwetschkeopsis Broth.
Theliaceae (Broth.) M. Fleisch. Plants terrestrial or on bases of trees;
stems monopodially branched; paraphyllia present; leaves imbricate,
little altered when moist, deltoid–ovate; costa single; laminal cells
short, stoutly unipapillose; alar cells differentiated; capsules
exserted, erect; exostome teeth white, smooth to papillose;
endostome strongly reduced; Thelia Sull.
Microtheciellaceae H. A. Mill. & A. J. Harr. Plants epiphytic; stems
monopodially branched?; costa single; laminal cells short, smooth;
alar cells weakly differentiated;, capsules short–exserted, erect;
exostome teeth truncate, reduced, weakly ornamented; endostome
rudimentary; Microtheciella Dixon
Sorapillaceae M. Fleisch. Leaves distichous and complanate; capsules
cladocarpous, immersed; peristome double, of 16 slender segments
and 32 stout exostome knobs, cilia absent; Sorapilla Spruce & Mitt.
Acknowledgments
We particularly appreciate the unpublished sequence data generously

provided by Cymon Cox. Thanks are also extended to Jon Shaw who also
provided unpublished sequence data and made helpful comments on the
manuscript and to Kevin Indoe who assisted with the digitization of the
photographic figures.
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Morphology and Classification of Mosses

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william r.

buck and bernard goffinet

Morphology and classiﬁcation of mosses

Bryophytes, in the broad sense, are the second largest phylum of

diversiﬁcation of mosses, as revealed by the extreme morphological diffe-

3.2 Synopsis of moss morphology

Here we present a general overview of the diversity of gametophytic and

Fig. 3.1. Polytrichum piliferum (Polytrichaceae), as an example of an acrocarpous moss.

Fig. 3.2. Helodium blandowii (Helodiaceae), as an example of a pleurocarpous moss.

not necessarily correlated with systematic position (see also La Farge-

Fig. 3.3. Brymela websteri (Pilotrichaceae), as an example of a moss. A. Aspect, showing

The external layer, or epidermis, is usually composed of relatively small,

structures are apparently physiologically active when the leaves are

surface. An extensive network of pores in the hyalocysts allows rapid

range of dispersal or abscission mechanisms for these gemmae, as well as

inﬂorescences. Perichaetial leaves are commonly used taxonomic charac-

3.2.2 Sporophyte and associated gametophytic tissues

division in particular, are central to most modern higher-level classiﬁca-

Fig. 3.4. Architecture of arthrodontous peristomes at maturity. One-eighth of the

During the ﬁnal stages of sporophyte maturation, additional material

Fig. 3.6. Schistidium heterophyllum (Grimmiaceae). Light micrograph of portion of the

two rows of teeth, an exostome and an endostome, while haplolepideous

Fig. 3.7. Tortula plinthobia (Pottiaceae). SEM of portion of the haplolepideous

Fig. 3.8. Pseudoscleropodium purum (Brachytheciaceae). SEM of portion of the

nodes or joints between two cells of a column (nodulose). The peristome

between obliquely rostrate opercula and cucullate calyptrae, and between

the Splachnaceae, differential thickening of walls is within individual

3.3 The classiﬁcation of mosses

Throughout the lifetimes of most living bryologists classiﬁcation systems

details of the peristome were considered as valuable in understanding

morphology and insertion, to structure of pseudoparaphyllia, to

3.4 A suggested classiﬁcation of the mosses

The following classiﬁcation is original in several aspects. Molecular data

adopted superorders, following Reveal’s (1998) suggestion to use the

CLASS: TAKAKIOPSIDA Leaves divided into terete ﬁlaments; capsules

Tetraphidaceae Schimp. Leaves unicostate; calyptrae small conic;

order: funariales Peristome lacking cilia, opposite, divisions in the IPL

Drummondiaceae (Vitt) Gofﬁnet, comb. nov. (Orthotrichaceae

Macrodictyum (Broth.) E. H. Hegew., Mesotus Mitt., Microcampylopus

Rhachitheciaceae H. Rob. Laminal cells rectangular in lower half,

Hymenostyliella E. B. Bartram, Hymenostylium Brid., Hyophila Brid.,

Bryobartramiaceae Sainsb. Plants very small, acrocarpous; calyptrae

Stoneobryum D. H. Norris & H. Rob., Ulota D. Mohr, Zygodon Hook. &

Schwägr., Bryum Hedw., Mielichhobryum J. P. Srivast., Mniobryoides

Calomniaceae Kindb. Plants small, arising from persistent protonema;

Hookeriaceae Schimp. Stems with central strand; pseudoparaphyllia

suborder: Ptychomniieae Plants usually robust and turgid, often

Platylomella A. L. Andrews, Sciaromiella Ochyra, Sciaromiopsis Broth.,

Dolichomitriopsis S. Okamura, Fabronidium Müll. Hal., Haplocladium

laminal cells elongate; alar cells often differentiated; setae not

Myriniaceae Schimp. Plants often epiphytic, small; stems

Fontinalaceae Schimp. Plants aquatic; stems sympodially branched;

Macrothamniella M. Fleisch., Mahua W. R. Buck, Microctenidium M.

(Broth.) Broth., Radulina W. R. Buck & B. C. Tan, Rhaphidostichum M.

endostome mostly rudimentary; spores often large and

absent; laminal cells linear, smooth, thick-walled and porose; alar

Lembophyllaceae Broth. Plants often turgid; stems monopodially

We particularly appreciate the unpublished sequence data generously

Dixon, H. N. (1932) Classiﬁcation of mosses. In Manual of Bryology, ed. F. Verdoorn, pp.

Koponen, A. (1990). Entomophily in the Splachnaceae. Botanical Journal of the Linnean

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