Moss Temporal range: Carboniferous[1] – recent PreЄ Є O S D C P T J K Pg N
"Muscinae" from Ernst Haeckel's Kunstformen der Natur, 1904
Scientific classification
Kingdom:	Plantae
Division:	Bryophyta Schimp.
Classes [2]
Takakiopsida Sphagnopsida Andreaeopsida Andreaeobryopsida Oedipodiopsida Polytrichopsida Tetraphidopsida Bryopsida

Mosses are small, soft plants that are typically 1–10 cm (0.4–4 in) tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems. At certain times mosses produce spore capsules which may appear as beak-like capsules borne aloft on thin stalks.

There are approximately 12,000 species of moss classified in the Bryophyta.^[2] The division Bryophyta formerly included not only mosses, but also liverworts and hornworts. These other two groups of bryophytes now are often placed in their own divisions.

Physical characteristics[link]

Description[link]

Botanically, mosses are bryophytes, or non-vascular plants. They differ from 'higher' plants by not having internal water-bearing vessels or veins, and no flowers and therefore no fruits, cones or seeds. They are small (a few centimeters tall) and herbaceous (nonwoody) and absorb water and nutrients through their leaves. Mosses have stems which may be simple or branched and upright or lax, simple leaves that often have midribs, roots (rhizoids) that anchor them to their substrate, and spore-bearing capsules on long stems. They harvest sunlight to create food through photosynthesis.^[3][4] Mosses do not absorb water or nutrients from their substrate through their roots, so while mosses often grow on trees, they are never parasitic on the tree.

In addition to lacking a vascular system, mosses have a gametophyte-dominant life cycle, i.e. the plant's cells are haploid for most of its life cycle.^[5] Sporophytes (i.e. the diploid body) are short-lived and dependent on the gametophyte. This is in contrast to the pattern exhibited by most "higher" plants and by most animals. In seed plants, for example, the haploid generation is represented by the pollen and the ovule, whilst the diploid generation is the familiar flowering plant.

They can be distinguished from the similar liverworts (Marchantiophyta or Hepaticae) by their multi-cellular rhizoids. Also, in most mosses, the spore-bearing capsule enlarges and matures after its stalk elongates, while in liverworts the capsule enlarges and matures before its stalk elongates.^[4] Other differences are not universal for all mosses and all liverworts, but the presence of clearly differentiated stem with simple-shaped, ribbed leaves, without deeply lobed or segmented leaves and not arranged in three ranks, all point to the plant being a moss.

Life cycle[link]

Most kinds of plants have two sets of chromosomes in their vegetative cells and are said to be diploid, i.e. each chromosome has a partner that contains the same, or similar, genetic information. By contrast, mosses and other bryophytes have only a single set of chromosomes and so are haploid (i.e. each chromosome exists in a unique copy within the cell). There are periods in the moss life cycle when they do have a double set of paired chromosomes, but this happens only during the sporophyte stage.

Life cycle of a typical moss (Polytrichum commune)

The life of a moss starts from a haploid spore. The spore germinates to produce a protonema (pl. protonemata), which is either a mass of thread-like filaments or thalloid (flat and thallus-like). Moss protonemata typically look like a thin green felt, and may grow on damp soil, tree bark, rocks, concrete, or almost any other reasonably stable surface. This is a transitory stage in the life of a moss, but from the protonema grows the gametophore ("gamete-bearer") that is structurally differentiated into stems and leaves. A single mat of protonemata may develop several gametophore shoots, resulting in a clump of moss.

From the tips of the gametophore stems or branches develop the sex organs of the mosses. The female organs are known as archegonia (sing. archegonium) and are protected by a group of modified leaves known as the perichaetum (plural, perichaeta). The archegonia are small flask-shaped clumps of cells with an open neck (venter) down which the male sperm swim. The male organs are known as antheridia (sing. antheridium) and are enclosed by modified leaves called the perigonium (pl. perigonia). The surrounding leaves in some mosses form a splash cup, allowing the sperm contained in the cup to be splashed to neighboring stalks by falling water droplets.

Mosses can be either dioicous (compare dioecious in seed plants) or monoicous (compare monoecious). In dioicous mosses, male and female sex organs are borne on different gametophyte plants. In monoicous (also called autoicous) mosses, both are borne on the same plant. In the presence of water, sperm from the antheridia swim to the archegonia and fertilisation occurs, leading to the production of a diploid sporophyte. The sperm of mosses is biflagellate, i.e. they have two flagellae that aid in propulsion. Since the sperm must swim to the archegonium, fertilisation cannot occur without water. After fertilisation, the immature sporophyte pushes its way out of the archegonial venter. It takes about a quarter to half a year for the sporophyte to mature. The sporophyte body comprises a long stalk, called a seta, and a capsule capped by a cap called the operculum. The capsule and operculum are in turn sheathed by a haploid calyptra which is the remains of the archegonial venter. The calyptra usually falls off when the capsule is mature. Within the capsule, spore-producing cells undergo meiosis to form haploid spores, upon which the cycle can start again. The mouth of the capsule is usually ringed by a set of teeth called peristome. This may be absent in some mosses. Most mosses rely on the wind to disperse the spores. In the genus Sphagnum the spores are projected about 10 to 20 cm off the ground by compressed air contained in the capsules; the spores are accelerated to about 36,000 times the earth's gravitational acceleration g.^[6][7]

In some mosses, e.g. Ulota phyllantha, green vegetative structures called gemmae are produced on leaves or branches, which can break off and form new plants without the need to go through the cycle of fertilization. This is a means of asexual reproduction, and the genetically identical units can lead to the formation of clonal populations.

Classification[link]

Traditionally, mosses were grouped with the liverworts and hornworts in the Division Bryophyta (bryophytes), within which the mosses made up the class Musci. This definition of Bryophyta, however, is paraphyletic and now tends to be split up into three divisions. In such a system, the Division Bryophyta contains exclusively mosses.

The mosses are grouped as a single division, now named Bryophyta, and divided into eight classes:

Moss in the Allegheny National Forest, Pennsylvania, USA.

Six of the eight classes contain only one or two genera each. Polytrichopsida includes 23 genera, and Bryopsida includes the majority of moss diversity with over 95% of moss species belonging to this class.

The Sphagnopsida, the peat-mosses, comprise the two living genera Ambuchanania and Sphagnum, as well as fossil taxa. However, the genus Sphagnum is a diverse, widespread, and economically important one. These large mosses form extensive acidic bogs in peat swamps. The leaves of Sphagnum have large dead cells alternating with living photosynthetic cells. The dead cells help to store water. Aside from this character, the unique branching, thallose (flat and expanded) protonema, and explosively rupturing sporangium place it apart from other mosses.

Andreaeopsida and Andreaeobryopsida are distinguished by the biseriate (two rows of cells) rhizoids, multiseriate (many rows of cells) protonema, and sporangium that splits along longitudinal lines. Most mosses have capsules that open at the top.

Polytrichopsida have leaves with sets of parallel lamellae, flaps of chloroplast-containing cells that look like the fins on a heat sink. These carry out photosynthesis and may help to conserve moisture by partially enclosing the gas exchange surfaces. The Polytrichopsida differ from other mosses in other details of their development and anatomy too, and can also become larger than most other mosses, with e.g. Polytrichum commune forming cushions up to 40 cm (16 in) high. The tallest land moss, a member of the Polytrichidae is probably Dawsonia superba, a native to New Zealand and other parts of Australasia.

They appear to be the closest living relatives of the vascular plants.

Red moss capsules, a winter native of the Yorkshire Dales moorland.

Geological history[link]

The fossil record of moss is sparse, due to their soft-walled and fragile nature. Unambiguous moss fossils have been recovered from as early as the Permian of Antarctica and Russia, and a case is put forwards for Carboniferous mosses.^[9] It has further been claimed that tube-like fossils from the Silurian are the macerated remains of moss calyptræ.^[10]

Habitat[link]

Dense moss colonies in a cool coastal forest

A closeup of moss on a rock

Young sporophytes of the common moss Tortula muralis (wall screw-moss)

Retaining wall covered in moss

The moss garden at the Bloedel Reserve, Bainbridge Island, Washington State.

Since mosses have no vascular system to carry water through the plant, they must have a damp environment in which to live, and a surrounding of liquid water to reproduce. And since mosses are photosynthetic, they require enough sun to conduct photosynthesis.^[5] Shade tolerance varies by species, just as it does with higher plants. In most areas, mosses grow chiefly in areas of dampness and shade, such as wooded areas and at the edges of streams; but they can grow anywhere in cool damp cloudy climates, and some species are adapted to sunny, seasonally dry areas like alpine rocks or stabilized sand dunes.

Choice of substrate varies by species as well. Moss species can be classed as growing on: rocks, exposed mineral soil, disturbed soils, acid soil, calcareous soil, cliff seeps and waterfall spray areas, streamsides, shaded humusy soil, downed logs, burnt stumps, tree trunk bases, upper tree trunks, and tree branches. Moss species growing on or under trees are often specific about the species of trees they grow on, such as preferring conifers to broadleaf trees, oaks to alders, or vice versa.^[4]

Mosses are also found in cracks between paving stones in damp city streets, and on roofs. Some species adapted to disturbed, sunny areas are well adapted to urban conditions and are commonly found in cities. Examples would be Rhytidiadelphus squarrosus, a garden weed in Vancouver and Seattle areas; Bryum argenteum, the cosmopolitan sidewalk moss, and Ceratodon purpureus, red roof moss, another cosmopolitan species. A few species are wholly aquatic, such as Fontinalis antipyretica, common water moss; and others such as Sphagnum inhabit bogs, marshes and very slow-moving waterways.^[4] Such aquatic or semi-aquatic mosses can greatly exceed the normal range of lengths seen in terrestrial mosses. Individual plants 20–30 cm (8–12 in) or more long are common in Sphagnum species for example.

Wherever they occur, mosses require high levels of moisture to survive because of the lack of a vascular system, and the need for liquid water to complete fertilisation. Many mosses can survive desiccation, sometimes for months, returning to life within a few hours of rehydration.^[5]

It is generally believed that in northern latitudes, the north side of trees and rocks will generally have more luxuriant moss growth on average than other sides.^[11] This is assumed to be because the sun on the south side creates a dry environment. South of the equator the reverse would be true. However, naturalists feel that mosses grow on the damper side of trees and rocks.^[3] In some cases, such as sunny climates in temperate northern latitudes, this will be the shaded north side of the tree or rock. On steep slopes it may be the uphill side. For mosses that grow on tree branches, this is generally the upper side of the branch on horizontally growing sections or near the crotch. In cool damp cloudy climates, all sides of tree trunks and rocks may be equally damp enough for mosses. And different species of mosses have different moisture and sun requirements so will grow on different sections of the same tree or rock.

Cultivation[link]

Moss is often considered a weed in grass lawns, but is deliberately encouraged to grow under aesthetic principles exemplified by Japanese gardening. In old temple gardens, moss can carpet a forest scene. Moss is thought to add a sense of calm, age, and stillness to a garden scene. Moss is also used in bonsai to cover the soil and enhance the impression of age.^[12] Rules of cultivation are not widely established. Moss collections are quite often begun using samples transplanted from the wild in a water-retaining bag. However, specific species of moss can be extremely difficult to maintain away from their natural sites with their unique requirements of combinations of light, humidity, substrate chemistry, shelter from wind, etc.

Growing moss from spores is even less controlled. Moss spores fall in a constant rain on exposed surfaces; those surfaces which are hospitable to a certain species of moss will typically be colonised by that moss within a few years of exposure to wind and rain. Materials which are porous and moisture retentive, such as brick, wood, and certain coarse concrete mixtures are hospitable to moss. Surfaces can also be prepared with acidic substances, including buttermilk, yogurt, urine, and gently puréed mixtures of moss samples, water and ericaceous compost.

In the cool cloudy damp Pacific Northwest, moss is sometimes allowed to grow naturally as a lawn substitute, one that needs little or no mowing, fertilizing or watering. In this case, grass is considered to be the weed.^[13] Landscapers in the Seattle area sometimes collect boulders and downed logs growing mosses for installation in gardens and landscapes. Woodland gardens in many parts of the world can include a carpet of natural mosses.^[5] The Bloedel Reserve on Bainbridge Island, Washington State, is famous for its moss garden. The moss garden was created by removing shrubby underbrush and herbaceous groundcovers, thinning trees, and allowing mosses to fill in naturally.^[14]

Mosses are sometimes used in green roofs. Advantages of mosses over higher plants in green roofs include reduced weight loads, increased water absorption, no fertilizer requirements, and high drought tolerance. Since mosses do not have true roots, they require less planting medium than higher plants with extensive root systems. With proper species selection for the local climate, mosses in green roofs require no irrigation once established and are low maintenance.^[15]

Inhibiting moss growth[link]

Moss can be a troublesome weed in containerized nursery operations and greenhouses.^[16] Vigorous moss growth can inhibit seedling emergence and penetration of water and fertilizer to the plant roots.

Moss growth can be inhibited by a number of methods:

• Decreasing availability of water through drainage.
• Increasing direct sunlight.
• Increasing number and resources available for competitive plants like grasses.
• Increasing the soil pH with the application of lime.
• Heavy traffic or manually disturbing the moss bed with a rake
• Application of chemicals such as ferrous sulfate (e.g. in lawns) or bleach (e.g. on solid surfaces).
• In containerized nursery operations, coarse mineral materials such as sand, gravel, and rock chips are used as a fast-draining top dressing in plant containers to discourage moss growth.