Classification Of Mosses 1h2457

CONTENT

CHAPTER I INTRODUCTION Mosses belong to the group of plants call Bryophytes which never produce flowers or seeds. They have a long history of evolution compared to flowering plants, with fossil record dated back 440 million years ago in the Silurian period. Most of the mosses are small, non-vascular, land plants, but some may be as large as 35cm tall and some are semi-aquatic. The mosses share a unique life cycle in having a first generation of dominant gametophyte. The gametophyte forms the green leafy structure we ordinarily associate with moss. Mosses are essentially land plants, and they depend on a film of water for the male cell (spermatozoid) to be transported to the female cell, both which are produced on the gametophyte. When conditions are right, the next generation, the sporophyte or spore-bearing structure is grown. The sporophyte is typically acapsule growing on the end of a stalk called the seta. The sporophyte has

no

chlorophyl

and

it

grows

parasitically

on

its gametophyte mother. The sporophyte dries out and releases spores that grow into

a

new

generation

of gametophytes,

if

they

germinate.

In general, when we are talking about mosses, we are generally referring to the gametophyte generation of the mosses' life cycle. Mosses do not have veins to transport water and food, instead, all parts of the plant are used to absord water and nutrients. Hence, mosses are found mostly at

places

where

there

are

a

consistent

supply

of

water.

Mosses

have rhizoids instead of roots, which are branched threads. While the rhizoids can absorb water, they are mainly used to anchor the plant.

CHAPTER II Classification of Mosses Division Anthocerotop hyta (Hornwortsh)

Class Anthocerotop sida Class Andreaeopsi da (Granite mosses)

Order Andreales Order Andreaebri ales Order Funariales

Superdivi sion Bryophyt es Kingdom Plantae

Division Bryophyta (Mosses)

Superdivis ion Pteridoph ytes Superdiviso n Spermatoph ytes Divison Hepatoph yta (Liverwort s)

Order Bryales Class Bryopsida (True mosses)

Class Sphagnop sida (Peat Mosses) Class Hepatopsi da

Order Buxbaumi ales Order Polytrical es Order Tetraphid ales Order Sphagnale s

Family Andreacea e Family Andreaebriace ae Family Funariace ae Family Bryaceae Family Buxbaumiac eae Family Polytricac eae Family Tetraphidac eae Family Sphagnac eae

ORDER SPHAGNALES The Sphagnales is an order of moss with only four living genera: Ambuchanania, Eosphagnum, Flatbergium, and Sphagnum

Figure: a clump of Sphagnum, peat moss FAMILY SPHAGNACEAE This family of this plant grows in swamps where the temperature is lower at high altitudes, such as in Dieng. Hyaline cell wall has a ring-shaped or spiral

thickening, hyaline cells that hold water. Peat moss leaves have a straight trunk and branches forming a rosette at the end. The sporogonium short-stemmed with foot haustorium (tool nutrient absorption) which later developed into pseudopodium. It spore capsule has closed but there are no peristome. Columella is hemispherical.

Figure: Sphagnum fimbriatum ANDREALES This order of plants reddish brown to greenish brown forming small tufts. The leaves falcate to secund-falcate, ovate-lanceolate to lanceolate. This order have the strong costa , laminal papillae.for the other characteristisc also include to dioicous, where the calyptras is tiny, campanulate-mitrate, seta lacking, sporophytes immersed to emergent on a gametophytic pseudopodium, peristome and operculum absent.

Figure.Andreaea nivalis Hook FAMILY ADREACEAE This plants of this family is reddish brown to greenish brown forming small tufts. Leaves falcate to secund-falcate, ovate-lanceolate to lanceolate, 0.6– 1.5 mm long, median cells quadrate to short-rectangular, 1+ stratose, papillose, 7–10

m in diameter, costa strong, percurrent, margins crenulate to strongly

denticulate. Dioicous, perichaetial leaves similar to the vegetative leaves but sometimes slightly larger, calyptra tiny, campanulate-mitrate, seta lacking, sporophytes immersed to emergent on a gametophytic pseudopodium, reddish brown to blackish up to 0.9 mm long when closed, peristome and operculum absent, opening by 4–6 longitudinal slits. Spores 18–40 m. Distinctive characters: (1) strong costa, (2) distinct crenulate to strongly denticulate leaf margins, (3) laminal papillae. Habitat Known from Greenland, Europe, Japan, and Russia. In the Pacific Northwest, found in Alaska, British Columbia, Washington, Oregon and California. Oregon Natural Heritage Information Center reports it from Clackamas and Lane Counties. In general Andreaea is often overlooked due to its

small size and is probably under collected because of its higher elevation tendencies. Habitat Associations: Andreaea nivalis forms reddish-brown mats on amp boulders in streamlet gullies, exposed rock outcrops, boulders next to melting snow, dry cliffs, sandy soil over boulders, and damp cliff faces in alpine to subalpine areas in the Pacific Northwest. Although referred to as a “granite moss” this genus often occurs on igneous rocks. It may form large mats or small patches and may or may not be abundant when found.

Figure. Andreae rupestris

ORDER ANDREBRIALES The distribution restricted to the northwestern part of Canada and adjacent Alaska, grows on calcareous rocks, in colder climates, contrasting with the acidic granite preference of Andreaea. ORDER FUNARIALES The order consists of two families, Disceliaceae , with a single species, and Funariaceae . Some species are annual and only grow for a few weeks when

conditions permit. An order of the true mosses (subclass Bryidae). The Funariales, often annuals or biennials and sometimes ephemeral, are for the most part characterized by a uniformity of gametophytic structure in contrast to a variability of sporophytic characters. Reduction in sporophytic characters is often associated with disturbed habitats and a shortened life cycle. FAMILY FUNARIACEAE This family of the plant minute to medium size which usually growing on soil. This also have light-green or yellow of color which gregarious of forming loose tufts. This have the long seta, slender is tough, reddish brown, stalk-like structure and the leaves, apex is acute to acuminate, costa single, smooth and rather thin-walled.

Funaria hygrometrica

Life Cycle This family have the same reproduction too vwith the common life cycle of the moses like this figure below.

Figure: The ife cycle of family funariaceae ORDER BRYALES This order is tree mosses which resemble small evergreen trees and are found in damp, shady places throughout the Northern Hemisphere. The branches clustered at the top of the shoot. The reddish-brown capsules (spore cases), borne on the female plant, have lids with long beaks and mature in the fall.

Bryum capillare

FAMILY BRYACEAE

Plants aquatic, forming loose floating mats, trailing from a single point of attachment, light to dark green. Stems usually elongate. The leaves unistratose, rather flaccid. Seta very short. Capsule immersed to emergent, oval cylindrical. Habitat Usually attached to submerged rocks and wood in slow to fast flowing streams or in ponds, but may be exposed for short periods when water levels drop. South African plants are believed to be introductions from Europe ORDER BUXBAUMIALES Buxbaumia (Bug moss, Bug-on-a-stick, Humpbacked elves, or Elf-cap moss) is a genus of twelve species of mosss (Bryophyta). It was named in 1801 by Johann Hedwigg to commemorate Johann Christian Buxbaumm, a German physician and botanist who discovered the moss in 1712 at the mouth of the Volga Riverr. The moss is microscopic for most of its existence, and plants are noticeable only after they begin to produce their reproductive structures. The asymmetrical spore capsule has a distinctive shape and structure, some features of which appear to be transitional from those in primitive mosses to most modern mosses.

Figure : Buxbaumia viridis

Plants of Buxbaumia have a much reduced gametophyte, bearing a sporophyte that is enormous by comparison. In most mosses, the gametophyte stage of the life cycle is both green and leafy, and is substantially larger than the spore-producing stage. Unlike these other mosses, the gametophyte of Buxbaumia is microscopic, colorless, stemless, and nearly leafless. It consists exclusively of thread-like protonemata for most of its existence, resembling a thin green-black felt on the surface where it grows.The plants are dioicous, with separate plants producing the male and female organs.Male plants develop only one microscopic leaf around each antheridium, and female plants produce just three or four tiny colorless leaves around each archegonium. Because of its small size, the gametophyte stage is not generally noticed until the stalked sporangium develops, and is locatable principally because the sporangium grows upon and above the tiny gametophyte. The extremely reduced state of Buxbaumia plants raises the question of how it makes or obtains sufficient nutrition for survival. In contrast to most mosses, Buxbaumia does not produce abundant chlorophyll and is saprophytic.It is possible that some of its nutritional needs are met by fungi that grow within the plant. The sporophyte at maturity is between 4 and 11 mm tall. The spore capsule is attached at the top of the stalk and is distinctive, being asymmetric in shape and oblique in attachment. As with most other Bryopsida, the opening through which the spores are released is surrounded by a double peristome (diplolepidious) formed from the cell walls of disintegrated cells. The exostome (outer row) consists of 16 short articulated "teeth". Unlike most other mosses, the endostome (inner row) does not divide into teeth, but rather is a continuous pleated membrane around the capsule opening.Only the genus Diphyscium has a similar peristome structure, although that genus has only 16 pleats in its endostome, in contrast to the 32 pleats in Buxbaumia. Diphyscium shares with Buxbaumia one other oddity of the sporophyte; the foot (stalk base) ramifies as a result of outgrowths, so much so that they may be mistaken for rhizoids.

Habitat Species of Buxbaumia may be found across much of the temperate to subarctic regions of the Northern Hemisphere, as well as cooler regions of Australia and New Zealand.

Figure : Sporophytes of Buxbaumia aphylla growing among other mosses. None of the visible leaves belong to Buxbaumia, which is a stemless and nearly leafless plant

The moss is an annual or biennial plant and grows in disturbed habitats or as a pioneer species. The plants grow on decaying wood, rock outcrops, or directly on the soil. They do not grow regularly or reliably at given locations, and frequently disappear from places where they have previously been found. Sporophyte stages begin their development in the autumn, and are green through the winter months. Spores are mature and ready for dispersal by the late spring or early summer. The spores are ejected from the capsule in puffs when raindrops fall upon the capsule's flattened top. The asymmetric sporophytes of Buxbaumia aphylla develop so that the opening is oriented towards the strongest source of light, usually towards the south.The species often grows together with the diminutive liverwort Cephaloziella, which forms a blackish crust that is easier to spot than Buxbaumia itself.

ORDER POLYTRICALES

This order have the best mosses vascular system to transport water from the ground leaves, stems cells having specialized in transporting water and plant epidermis waterproofing, so that unlike typical mosses where water is transported by capillarity through the epidermis, the transport in these species is internal. FAMILY POLYTRICACEAE The Polytrichaceae is a common family of mosses. of this family tend to be larger than other mosses with a thickened central stem and a rhizome. The leaves have a midrib that bears lamellae on the upper surface. Species in this group are dioicous. Another characteristic that identifies them is that they have from 32 to 64 peristome teeth in their sporangium. A thickened central stem and a rhizome. The leaves have a midrib that bears lamellae on the upper surface. In this family the hard leaves with lamellar many expansions will make the sheet retains water surface and increase photosynthesis. Habitat This moss live wildly and become the common hair cap moss that is easy to find. Preferring to live in lightly shaded areas with moist slightly acidic soil, it can also survive in areas of full sunlight provided the soil is moist. The common hair cap moss can also grow in areas of poor soil and slow drainage Reproduction The female shoots develop the eggs and the male shoots develop the sperm. In the spring, raindrops splash the sperm from the male shoots to the female shoots where they then travel into the egg. During the summer spores are released and carried by the wind. When the spore reaches a habitat it can survive in, it germinates and the process starts over again

One of the example of this family is Polytrichum commune

Kingdom : Plantae Phylum : Bryophyta

Genus

:Polytrichum

Species

: Polytrichum commune

Life Cycle of Polytricaceae

Class

: Bryopsida

Order

: Polytrichales

Family

: Polytrichaceae

ORDER TETRAPHIDALES This order has single family Tetraphidaceae represented by two genus Tetraphis

and

Tetradontium.

Tetraphis

is

northen

Hemisphere

genus,

inhabitingconiferus forests. The small plants grow on decomposing wood, prat banks and rarely on sandstone. FAMILY TETRAPHIACEAE The leaves are small but those on the upper stem are big bright green, ovate and costate.This plants are monoceous, the archegonia and antheredia are on terminal

separate

branches.This

order

has

2

genus:

Tetraphis

and

Tetrodontium.This moss can live in moist place that have aciditic or neutral PH. One of the example of tetraphiaceae is Tetraphis pellucida

Kingdom: Tetraphis Division: pellucida

Plantae

Bryophyta The

Class:

Bryopsida

morphology

Tetraphis

pellucida

of are

Stem are not rhizomes, costa slender, capsule long-

Order:

Tetraphidales

Family:

Tetraphidales

Genus:

Tetraphis

Species:

Tetraphis pellucida

exserted and leave small.

Division Anthocerotop hyta (Hornwortsh)

Class Anthocerotop sida Class Andreaeop sida (Granite mosses)

Superdivi sion Bryophyte s Kingdom Plantae

Division Bryophyta (Mosses)

Superdivis ion Pteridoph ytes Superdiviso n Spermatoph ytes

Class Polytrichopsi da

Order Diphyscia ales

Family Diphysciac eae

Order Buxbaumi ales

Family Buxbaumiace ae

Order Polytricale s

Family Polytricac eae

Order Tetraphida les

Family Tetraphidac eae

Class Andreaeobryo psida Divison Hepatoph yta (Liverwort s)

Class Hepatopsida

Actually we got another source about the mosses classification, here below the graph.

Andreaeopsida – the granite mosses This is a small, cool-climate class of siliceous-rockdwelling mosses (Schofield 1985), again with only one genus, but with approximately 100 species. They are typically blackish or reddish, brittle, and short (Figure 1). One can recognize them by rubbing one's hand across them and discovering small fragments stuck to the hand. The leaves are but one cell thick, but some species have a multiple cell thickness in the center, forming a costa. The arrangement of leaves is multi-ranked. Unlike most mosses, they have a thalloid protonema. The capsule is reminiscent of liverworts, opening in four valves, but having the tips remaining attached to each other, making it look like those paper lanterns we made as children for Halloween (Figure 2). Unlike the liverworts, it lacks elaters. And unlike most liverworts and Bryopsida, it lacks a seta and has a gametophyte

pseudopodium, a character in common with Sphagnopsida, a stalk produced at capsule maturity from the gametophyte tissue. Of ecological significance, it is autoicous (having male and female reproductive organs in separate clusters on the same plant). This ensures there will be others around to accomplish fertilization.

Figure 2. Andreaea rupestris, Class Andreaeopsida, gametophyte with sporophyte showing four valves of capsule and pseudopodium of gametophyte.

Andreaeobryopsida This class likewise is comprised of a single genus, Andreaeobryum (Figure 3), which has been considered by most to belong to the Andreaeopsida, but recently separated in the treatment by Buck and Goffinet (2000). It differs in being dioicous (having male and female reproductive organs on separate plants) and possessing a seta. Its calyptra is larger, covering the capsule, and the capsule is valvate, but unlike the Andreaeopsida, the apex erodes, so the valves are free, not ed at the apex. The distribution is narrow, restricted to the northwestern part of Canada and adjacent Alaska, where it grows on calcareous rocks, contrasting with the acidic granite preference of Andreaea.

Figure 3. Andreaeobryum macrosporum with valvate capsules. Photo from Biology 321 Course Website, http://www.botany.ubc.ca/bryophyte/LAB6b.htm. Polytrichopsida With bryophytes, the determination of primitive or advanced often depends on the generation being examined. The gametophyte may have changed considerably while some set of characters of the sporophyte remained constant. And of course, the reverse can be true. The dioicous condition (male and female reproductive organs on separate plants) that characterizes Polytrichopsida is considered to be primitive (Longton & Schuster 1983), with the monoicous condition (male and female reproductive organs on the same plant) that is so frequent in Bryopsida typically being derived by doubling of the chromosome number. Likewise, nematodontous peristome teeth (having evenly thickened walls and whole dead cells lacking eroded walls, Figure 4) of Polytrichopsida would seem to be an earlier development than the arthrodontous condition of Bryopsida. All of the class possess an elongate sporophyte seta, ing an operculate peristomate capsule, and a columnar columella, characters that are more advanced than in Sphagnopsida but typical in Bryopsida. Spores are produced by meiosis in a single event in sporogenous tissue that surrounds the columella (Figure 5).

Figure 4. Nematodontous peristome teeth of Tetraphis pellucida (Polytrichopsida). Note the separation at the tips. Photo from Biology 321 Course Website, www.botany.ubc.ca/ bryophyte/LAB6b.htm.

Figure 5. Cross section of immature Polytrichum capsule showing sporogenous tissue. Photo by Janice Glime.

The gametophyte is often very specialized, being characterized by stems

with

a

central

strand,

reaching its peak in Polytrichaceae, with the presence of hydroids (water-conducting

cells)

and

leptoids (sugar-conducting cells). The leaves of the class are all costate

(having

a

midrib-like

structure).

Figure 6. Longitudinal section of Polytrichum capsule. Photo by Janice Glime.

Figure 7. Cross section of a Polytrichum stem showing green hydroids in center and larger leptoids surrounding them. Photo by Izawa Kawai.

Polytrichaceae In many ways, this family looks like a tracheophyte wanna-be. It attains a greater height than the typical moss and can even stand alone to nearly half a meter in the case of Dawsonia superba. Polytrichum commune likewise attains similar heights, but only with the of other individuals, forming a hummock.

Figure 8. Dawsonia superba from New South Wales, Australia. Photo by Janice Glime. The Polytrichaceae lead the way to complexity with their unusual leaf structure, possessing vertical lamellae (vertical tiers of cells like the pages of an open book; (Figure 9) that provide an interior somewhat resembling that of a maple leaf. In fact, in the genus Polytrichum, some have the outer portion of the blade folded over the lamellae, creating an internal chamber resembling palisade mesophyll surrounded with epidermis. The cuticle (in this case, a waxy, water-repellant covering on the outer surface of the leaf; Proctor 1979) of Polytrichum is more developed than in most other bryophytes, and Polytrichum seems to repel water from its leaves rather than to absorb it (Figure 10), a phenomenon that may prevent the spaces among the lamellae from flooding that would block access of CO2 to the chloroplasts within. Its rhizoids function not only for anchorage, but also seem to facilitate external water movement. Figure 9. Leaf cross sections of Polytrichopsida. Left: Stained section of Polytrichum leaf showing vertical lamellae. Right: Hand section of Polytrichastrum alpinum leaf showing lamellae with papillose terminal cells. Photos by Janice Glime.

Figure 10. Polytrichum juniperinum with waxy leaves that roll over the lamellae. Photo by Janice Glime. In some mosses, like Polytrichum, the antheridia are in splash cups or platforms (rosette of leaves from which reproductive units such as sperm, gemmae, or spores can be splashed by raindrops; Figure 11), and when the sperm (male reproductive cells; male gametes) are mature, the antheridium swells and bursts during a rainy period. The bases of the antheridia, in taxa such as Polytrichum and Atrichum, collect fluid between the sperm tissue and the antheridial jacket (Bold et al. 1987). When the cells at the tip of the sterile jacket open, the antheridial jacket contracts. At this time, the fluid at the bottom acts as a hydraulic ram and forces the sperm out of the antheridium. Once in the open water of the splash cup, the sperm are splashed from the cup. Hopefully, some of these sperm will be splashed near the tip of a female plant (Figure 12) and will begin swimming toward the archegonium. But it appears that the sperm of Polytrichum

commune, and perhaps others, may have some help in this process from another source (Harvey-Gibson & Miller Brown 1927). A variety of invertebrates visit the male splash cups once they are fertile and get the mucilage with sperm stuck on their bodies. While visiting the plants, the insects lap up the mucilage and lick the saline crystals that form on the margins of the perichaetial leaves. The same insects, bodies and limbs smeared with mucilage in which sperms were abundant and motile, likewise appear on female plants. Now, can someone show whether the red color of splash cups (Figure 11) in several of this family have the ability to attract any dispersal agents?

Figure 11. Male plants of Polytrichum juniperinum withantheridial splash cups. Photo by Janice Glime.

Figure 12. Female plants of Polytrichum ohioense showing the tight leaves at the apex where archegonia are housed. To th right of the female plants, the yellow swollen tips are unopened antheridial splash cups. Photo by Janice Glime.

After fertilization, the zygote to

divides

form

embryo

an

within

the archegonium. Eventually this sporophyte tissue forms a foot, seta, and capsule.

The

capsule develops within the calyptra which is the old archegonium. The calyptra is essential for normal development in most mosses, and a split on one side can cause asymmetrical development. In the case of Polytrichum, the calyptra is very hairy, earning the moss the name of hairy cap moss or goldilocks moss. Eventually the calyptra is shed, exposing the capsule. Then the operculum (lid) must come off to permit spore dispersal. In this family the capsule has 64 short teeth ed by a membrane (epiphragm) that covers the capsule like skin on a drum. These small spaces permit spores to escape the capsule a few at a time, providing maximum chances for some escaping under the right conditions for dispersal and establishment.

Atrichum (Polytrichaceae) life cycle stages. Left: Atrichum undulatum leaves. Middle: Atrichum undulatum antheridial splash cups. Right: Atrichum angustatum capsules. Photos by Janice Glime. Tetraphidaceae Tetraphis , also in the Polytrichopsida, looks more like a typical moss than do other Polytrichopsida, with thin, 1-cell-thick leaves and a costa. Tetraphis is unique among mosses in having gemmae arranged in splash cups at the tips of the stems when sexual reproduction is not in season, arguably a primitive remnant. These gemmae are asexual bits of plant material that can grow into a new plant. Its most unusual character is that its protonemata are not threads, but rather flaps . Antheridia are borne terminally on the leafy plants ,as are the archegonia. The capsule has only

four

long, uned,

nematodontous Teeth. Protonemal flaps of Tetraphis pellucida. Photos from Biology 321 CourseWebsite,www.botany.ubc.ca/bryophyte/LAB6b.htm

Buxbaumiaceae – Bug on a Stick Buxbaumia (Figure 23) is one of the strangest of all mosses. It lacks any leafy stem at all. Its archegonia and antheridia arise directly from the protonema. Hence, its capsules arise directly from this persistent protonema . Its capsules, although possessing teeth, more typically split across their broad, flattened surface, hence exposing the spores. The capsule interior is chambered and spongy, somewhat like a spongy mesophyll of Magnoliophyta. It typically occurs with tiny, black leafy liverworts such as Cephalozia.

Unopened capsule of Buxbaumia aphylla,illustrating the flat side with a beaked operculum that has earned it the common names of bug-on-a-stick and Aladdin's lamp moss. The sporophyte originates from a protonema with no leafy gametophyte. Photo by Michael Lüth. Diphysciaceae Perhaps related to Buxbaumia, the moss Diphyscium has a capsule of similar shape and lacks a leafy female stem, but the male plant of this genus has large, strap-shaped leaves and leads an independent and separate existence.

Upper: Diphyscium foliosum female plants with young sessile capsules among male plants. Photo by Janice Glime.

Lower: Mature female Diphyscium foliosum plants with capsules showing peristome teeth. Photo by Michael Lüth.

CHAPTER III CONCLUTION Bryophyta can be considered to have six classes: Takakiopsida, Sphagnopsida, Andreaeopsida, Andreaeobryopsida, Polytrichopsida, and Bryopsida, differing most consistently in capsule structure. Gametophores of Andreaeopsida, Andreaeobryopsida, and Polytrichopsida produce archegonia and/or antheridia at the apex and the embryo develops within the archegonium. Sporophytes remain attached to the gametophyte and produce spores by meiosis. These classes, and all Bryophyta, produce spores from the sporophyte only once. Takakiopsida, Andreaeopsida, and Andreaeobryopsida have capsules that split into valves, but lack elaters. Sphagnopsida lacks valves and has an operculum that is shed at dispersal time, but lacks peristome teeth. In capsules of Polytrichopsida and Bryopsida, an operculum usually covers peristome teeth that

often aid dispersal, contrasting with liverworts wherein the capsule splits into four valves with elaters that possibly facilitate spore movement. Polytrichopsida have nematodontous peristome teeth; Bryopsida have arthrodontous peristome teeth. All other classes of Bryobiotina lack peristomes. Andreaeobryopsida is dioicous (two sexes on separate plants) and possesses a seta (stalk of capsule), whereas Andreaeopsida is monoicous (both sexes on same plant) and lacks a seta.

REFERENCES

Bold, H. C., Alexopoulos, C. J., and Delevoryas, T. 1987. Morphology of Plants and Fungi. Harper & Row, Publishers, Inc., New York, NY. 912 pp. Buck, W. R. and Goffinet, B. 2000. Morphology and classification of mosses. In: Shaw, J. A. and Goffinet, B. (eds.). Bryophyte Biology. Cambridge University Press, pp. 71-123. Harvey-Gibson, R. J. and Miller Brown, D. 1927. Fertilization of Bryophyta. Preliminary note. Ann. Bot. 49: 190-191. Longton, R. E. and Schuster, R. M. 1983. Reproductive biology. In: Schuster, R. M. (ed.). New Manual of Bryology, Vol. 1, pp. 386-462. Proctor, M. C. F. 1979. Surface wax on the leaves of some mosses. J. Bryol. 10: 531-538.

Schofield, W. B. 1985. Introduction to Bryology. Macmillan Publishing Co., New York, 431 pp.