MTU.edu: Chapter 2-7 Bryopsida

CHAPTER 2-7 

BRYOPSIDA 

Figure 1. Aulacomnium androgynum with asexual gemmae on a modified stem tip. Photo by Michael Lüth. 

Bryopsida Definition 

By far the largest class of Bryophyta (sensu stricto) 

(84% of families) (Goffinet et al. 2001) and ~98% of the 

species, this class is unquestionably the most diverse. 

Their evolution by both advancement and reduction makes 

circumscription difficult, with nearly every character 

having exceptions. It appears that the only unique and 

consistent character among the Bryopsida is its peculiar 

peristome of arthrodontous teeth (the lateral walls of the 

peristome teeth are eroded and have uneven thickenings; 

Figure 2). 

This arrangement of teeth has implications for 

dispersal – the teeth form compartments in which spores 

are trapped. The outer surface is hydrophilic (water 

loving, hence attracting moisture) whereas the inner layer 

has little or no affinity for water (Crum 2001), causing the 

teeth to bend and twist as moisture conditions change. 

Whether this aids or hinders dispersal, and under what 

conditions, is an untested question. Yet even this character 

does not hold for some taxa; some taxa lack a peristome. 

And all other characters, it would seem, require the 

adjectives of most or usually. 

Figure 2. Electron micrograph of the arthrodontous 

peristome teeth of the moss Eurhynchium praelongum. Photo 

from Biology 321 Course Website, 

http://www.botany.ubc.ca/bryophyte/LAB6b.htm. 

45

46 Chapter 2-7: Bryopsida 

Spore Production and Protonemata 

As in all bryophytes, the spores are produced within 

the capsule by meiosis. In the Bryopsida, once germinated 

(Figure 3), they produce a filamentous protonema (first 

thread) that does not develop into a thalloid body (Figure 

4). This germination process can be rapid (1-3 days in 

Funaria hygrometrica) or lengthy, involving a long 

dormancy period. 

Figure 3. Germinating spore of Fontinalis squamosa. Photo 

by Janice Glime. 

Figure 4. Protonemata among leafy plants of Plagiomnium. 

Photo by Janice Glime. 

Many mosses differentiate their protonemata into 

chloronema and caulonema (Figure 5, Figure 6). The 

chloronema, meaning light green thread or chlorophyll 

thread, is the first part of the protonema to form when the 

spore germinates. The caulonema, meaning stem thread, is 

the portion that develops later, but not in all mosses, and 

that gives rise to the upright gametophores, or leafy plants. 

The caulonema differs from the younger parts of the 

protonema, the chloronema, in having longer cells with 

slanting cross walls, usually brownish cell walls, and fewer, 

less evenly distributed, smaller spindle-shaped chloroplasts. 

The chloronema exhibits irregular branching, whereas the 

caulonema exhibits regular branching. 

Figure 5. Protonema of moss such as Funaria hygrometrica 

with differentiated caulonema and chloronema. Drawing by Noris 

Salazar Allen. 

Figure 6. Protonema of Funaria hygrometrica showing 

chloronema (short cells with perpendicular walls and dense 

chloroplasts) and caulonema (long cells with diagonal cross walls 

and more dispersed chloroplasts). Photo by Janice Glime. 

Gametophore Buds 

As the protonema continues to develop and produce 

buds (Figure 7, Figure 8), the mosses and liverworts again 

differ. In liverworts, the bud is produced by the apical cell, 

hence ending further growth of the protonema and 

accounting for its single gametophore. In mosses, on the 

other hand, the bud originates from a cell behind the apical 

cell, hence permitting the apical cell to continue to divide 

and the protonema to continue to grow. The result is that 

moss protonemata produce many buds and upright plants 

(Figure 9). This provides the possibility for somatic 

mutations to arise, affording genetic variation among the 

leafy plants. 

Figure 7. Protonema and young developing bud of the moss 

Funaria hygrometrica. Photo by Martin Bopp. 

As the bud develops, rhizoids (Figure 25) form, 

functioning largely in anchorage, but at least in some 

mosses, also functioning in moving water and nutrients 

from substrate to moss. This may be especially important 

as the atmosphere dries and the rhizoids help to maintain a 

humid substrate.

Figure 8. Moss protonema with developed bud. Photo by 

Janice Glime. 

Figure 9. Leafy buds on the protonemata of Funaria 

hygrometrica forming a doughnut shape. Each of these circles of 

buds is the result of one spore. The hole in the middle is the area 

where the protonemata is in the chloronema stage and does not 

produce buds. Photo by Janice Glime. 

Gametophores 

The bud develops into the upright (or horizontal) 

gametophore. These plants are leafy haploid (1n) plants; 

thus, they are the dominant gametophyte generation of 

the life cycle. The stem may have a central strand (Figure 

10), or lack it (Figure 11); this strand may or may not have 

hydroids. 

Their leaves, more accurately known as phyllids (but 

rarely called that), are usually in more than three rows, 

but there are exceptions with two or three rows. Typically 

they are one cell thick, but there are modifications on this 

scheme that are expressed in some mosses by leaves folded 

over on themselves, creating a pocket in the genus 

Fissidens (Figure 12), or alternating hyaline (colorless) and 

photosynthetic layers as in Leucobryum, or just multiple 

layers of tissue, sometimes in patches. Some may have 

borders which likewise can be one or more layers thick. 

These leaves often have a multi-layered costa (Figure 13) 

in the center, or double, or even triple costa. The costa 

itself (Figure 25) consists of long, narrow cells that offer 

support and seem to function in moving water more quickly 

than their wider and often shorter neighboring cells. 

Chapter 2-7: Bryopsida 47 

Figure 10. Stem cross section of Rhizogonium illustrating 

central strand of hydroids. Photo by Isawa Kawai. 

Figure 11. Cross section of stem of the brook moss 

Fontinalis dalecarlica showing absence of central strand and 

conducting tissues. Photo by Janice Glime. 

Figure 12. Pockets in leaf of Fissidens arnoldii. Photo by 

Michael Lüth. 

Figure 13. Cross section of Bryopsida leaf showing one cell 

thick lamina (blade) portion and thickened costa. Photo by Janice 

Glime.


Location of Sex Organs 

Based on the branching patterns and location of sexual 

organs, the Bryopsida have traditionally been divided into 

two major groups, although there are good arguments for 

additional groupings. The acrocarpous mosses (Figure 

14) are generally those upright mosses with terminal 

sporangia. They usually are unbranched or sparsely 

branched. Pleurocarpous mosses (Figure 15), by contrast, 

produce their sporangia on short, specialized lateral 

branches or buds and typically are prostrate, forming freely 

branched mats. The truly pleurocarpous mosses appear to 

represent a single monophyletic clade (Buck & Goffinet 

2000; Buck et al. 2000a, b; Cox et al. 2000) and may be an 

adaptation to forming mats of continuous growth in mesic 

conditions (Vitt 1984). Those mosses that bear 

sporophytes terminally on short, lateral branches form a 

special category of pleurocarpous mosses termed 

cladocarpous. The branching patterns and positions of 

sporangia determine not only the growth form, but also 

influence success of fertilization, availability of water, and 

ability to spread horizontally across a substrate. 

Figure 14. Barbula unguiculata, an acrocarpous moss. 

Setae originate at the apex of the previous year's growth. Photo 

by Michael Lüth. 

Figure 15. Pleuroziopsis ruthenica, a pleurocarpous moss. 

Photo by Janice Glime. 

The upright or sprawling stems of the gametophyte 

produce antheridia (sperm-containers) and archegonia 

(egg-containers). In mosses, antheridia and archegonia 

may be located at the end of the main stem, at the ends of 

lateral branches, or along the main stem, either at the ends 

of very short branches or nearly sessile (Figure 26). Often 

the chloroplasts of the antheridial jacket cells are converted 

into chromoplasts as the antheridia mature, causing the 

characteristic red-orange color (Bold et al. 1987; see Figure 

16). 

Sperm Dispersal 

The dispersal of the sperm from the antheridium is an 

interesting phenomenon. In Mnium hornum, within about 

four minutes of placing water into an antheridial cup, 

dehiscence will occur (Muggoch & Walton 1942). The 

spermatocytes (cells in which sperm have differentiated) 

emerge in a banana-shaped package into the water 

surrounding the antheridium, usually within 4-10 minutes. 

Then, when (or if) that package connects with the water-air 

interface, the sperm spread apart rapidly to form a surface 

layer of regularly spaced sperm. This suggests that some 

substance with a low surface tension might be present in 

the sperm package because the mass spreads much like an 

oil spill. The behavior suggests that there is a small 

amount of fat present in the sperm mass. Muggoch and 

Walton considered this to be a widespread phenomenon, 

perhaps true of all mosses, and that it was important in 

permitting insects to carry sperm to female plants. 

However, there seem to be few observations of such insect 

dispersal except in Polytrichum (Class Polytrichopsida) and 

Bryum (or Rosulabryum) capillare (Bryopsida; Figure 16). 

Figure 16. Bryum capillare showing antheridial head of 

male plants. Photo by Michael Lüth. 

It appears that Bryum capillare may indeed be 

fertilized, at least some of the time, by animals. When 

covered by a fine net to discourage winged insects and 

other creatures, females were not fertilized, but when the 

net was removed, fertilization occurred 2 m(!) from the 

nearest males (Gayat 1897). However, it is difficult to rule 

out the possibility of raindrops in this case, or even 

squirrels, for that matter. Raindrops are likely to trap the 

mucilage with its sperm load in the tiny capillary spaces of 

the net. The success of fertilization would depend on the 

success of these drops getting bounced from one plant to 

another, and that bounce would surely be inhibited by such 

a filter to diminish the impact and retain the mucilage. 

Observations on Bryum argenteum are more 

conclusive. Cronberg et al. (2006), in an experiment in

which male and female plants were separated by 0, 2, and 4 

cm, demonstrated that help from such agents as 

invertebrates are essential. These treatment distances were 

combined either with no animals, or with mites (Acarina: 

Scutovertex minutus) or springtails (Collembola: Isotoma 

caerulea, Figure 17) (Cronberg et al. 2006; Milius 2006). 

After three months, those females in contact with male 

plants (0 cm) produced sporophytes. Those without this 

contact (2 or 4 cm) and without either animal group 

produced no sporophytes. But those housed with 

springtails or with mites produced numerous sporophytes, 

with springtails being the more effective conveyor. 

Springtails are more mobile than mites, and in this 

experiment, more sporophytes were produced at greater 

distances when springtails were available as dispersal 

agents. 

Figure 17. Isotoma caerulea, a springtail that is instrumental 

in fertilizing Bryum argenteum. Photo by Katrina Hedlund. 

But how do these springtails find the mosses? Flowers 

provide odors and colors to attract their pollinators. It 

appears that these mosses also have a way to attract their 

dispersal agents. When springtails and mites were given 

choices of plants with mature gametangia vs those that 

were sterile, fertile plants were chosen over non-fertile ones 

about five times as often (Beckman 2006) in the cases of 

both males and females and by both organisms. Cronberg 

et al. (2006) suggest that fertile plants may attract the 

invertebrates with sucrose (Pfeffer 1884), starch, fatty 

acids, and/or mucilage (Harvey-Gibson & Miller-Brown 

1927; Paolillo 1979; Renzaglia & Garbary 2001). 

Anderson (2002) managed to catch the dispersal of 

Plagiomnium affine on video to see the effectiveness of the 

splash platform of that moss. Although many drops will 

miss the tiny platform completely, a few manage full hits. 

Impact causes a "crown" of water to form, like dropping a 

rock into a lake. The capillary spaces between the 

antheridia and adjoining paraphyses (sing. paraphysis: 

sterile filaments located among reproductive organs; Figure 

18, Figure 26, Figure 27) fill with water. The impact of the 

drop causes the swollen antheridia to burst, releasing the 

swimming sperm. For the splash to be effective in making 

the crown, the diameter of the drop should be 1 mm or less, 

a common size in most rain showers. The rim of the crown 

has small droplets that are propelled away by the action. 

Since these droplets include water from within the splash 

platform, they also contain the sperm and thus propel them 

away from the plant. These droplets can travel 100 mm or 

more and manage to fertilize most of the females within 80 


mm. The dioicous liverwort Marchantia has a splash 

platform that performs a similar function. 

Splash cups and platforms seem to be rare in 

monoicous taxa (exceptions include species of 

Brachymenium and Rosulabryum per John Spence), 

suggesting fertilization is accomplished with close 

neighbors. For most Bryopsida, however, there is no 

antheridial splash cup or platform, so seemingly sperm 

must swim all the way. However, other things can create 

splash. Jonathan Shaw (pers. comm.) has considered that 

F. hygrometrica has wide-spreading bracts surrounding the 

antheridia and the flexible nature of these bracts permits 

them to bend back and create an effective cup from which 

sperm in that species might be splashed. Angela Newton 

(pers. comm.) has suggested that platform surfaces among 

the more dendroid and shelf-forming taxa could be viewed 

as water-trapping mechanisms that would promote surface 

flow and dripping to the next level down as a mode of 

transporting sperm between individual plants or parts of 

plants. One complication in this arrangement is that the 

complex texture would act to trap water drops rather than 

encouraging them to splash out and away. However, in 

some of the plants with large smooth leaves, these leaves 

might act as springboards, but she considered that in such a 

case the water drops would be unlikely to carry sperm, 

although they might carry the smaller kinds of vegetative 

propagules. Nevertheless, sperm that had gotten as far as a 

leaf might benefit from this splash as well. 

Figure 18. Mature antheridia and paraphyses of the moss 

Rhizomnium. Photo by Janice Glime. 

Whereas flowering plants frequently rely on animals, 

especially insects, to transport their male gametophytes, 

and ultimately the sperm, to the female reproductive organ, 

this seems rarely to be the case in bryophytes. 

Surprisingly, it appears that the only documented case of 

such animal transport of sperm is in Polytrichum commune 

(Polytrichopsida), which has well-developed splash cups 

for the purpose of sperm dispersal. Nevertheless, it was in 

this species that Harvey-Gibson and Miller-Brown (1927) 

found motile sperm on the bodies of small arthropods 

(flies, leafhoppers, mites, spiders, and springtails) on both 

male and female reproductive inflorescences. Schofield 

(1985) suggests that mucilage produced in both the 

perigonia (modified leaves enclosing male reproductive 

structures) and perichaetia (modified leaves enclosing


female reproductive structures) sometimes attracts 

invertebrates. 

One might expect that many antheridia burst as they 

and their surrounding paraphyses swell from a desiccated 

state to a hydrated state during early minutes of a 

precipitation event. Could it be that the same external 

capillary forces that carry water rapidly to other parts of the 

plant could move sperm, thus reducing the energy 

requirements for getting these tiny cells to their 

destinations? Or are these forces to be reckoned with, 

forcing the sperm to swim against a current? 

If animal dispersal is so rare, then how, in this vast 

world, does an unintelligent sperm find an archegonium 

(Figure 19) and an egg? Fortunately for the moss, the 

archegonium at this time has dissolved the neck canal cells 

(entry canal through neck to egg in base of archegonium; 

Figure 26) leading down to the egg, and the resulting liquid 

provides a chemical attractant for the sperm. Meanwhile, 

the egg exudes mucilage into the cavity of the venter (Lal 

et al. 1982). When the canal opens, the liquid exudes from 

the opening of the neck, creating a chemical gradient. The 

sperm follows the concentration gradient toward the 

archegonium and finally swims down the neck canal of the 

archegonium to the egg. The exact nature of this liquid is 

unknown, but it seems that sugars and sometimes boron are 

necessary. It seems also likely that something specific, 

perhaps a protein, might guide the sperm to the correct 

species. Otherwise, it would seem that in spring, when so 

many species are producing sexual structures, some of 

these sperm would find their way into the wrong 

archegonium – or perhaps they do! 

Figure 19. Archegonia of the moss Fontinalis duriaei. 

Photo by Janice Glime 

Embryo Development 

When a sperm reaches and fertilizes an egg, the 

resulting diploid (having two sets of chromosomes; 2n) 

zygote begins dividing by mitosis to form an embryo that 

starts to stretch the archegonium (Figure 20). But the 

archegonium cannot stretch indefinitely, and as the embryo 

gets larger, the archegonium finally tears. Here, mosses 

and liverworts differ. In most mosses, part of the 

archegonium remains perched on top of the developing 

embryo (young sporophyte). This separated piece of 

archegonium is the cap you often see on top of the capsule 

and is now called a calyptra (Figure 26). So the calyptra is 

a 1n covering over the 2n capsule. 

The emerging embryo grows into the sporophyte of the 

moss. The mature sporophyte has a capsule and stalk 

(seta), with a foot embedded into the gametophyte tissue 

(Figure 21). Meiosis occurs in the mature capsule, 

producing haploid (1n) spores, as in all plants. Note that 

this is a major difference from meiosis in animals, which 

results in gametes. These spores are dispersed from the 

capsule by wind (or in a few cases – Splachnaceae – by 

insects) and grow into new gametophytes. 

Figure 20. Development of the calyptra of a moss. a. egg 

in archegonium, with neck canal cells not yet disintegrated. b. 

archegonium after fertilization and early development of embryo, 

showing elongation of the archegonium as embryo grows. c. 

elongated seta with calyptra perched on top of it before capsule 

has developed. d. mature capsule with calyptra and fully 

elongated seta. c & d indicate the remains of the venter of the 

archegonium at the base of the sporophyte. Drawings by Janice 

Glime. 

The calyptra that covers the capsule of mosses most 

likely plays multiple roles. We know that in many species, 

normal development ceases if the calyptra is removed 

(Paolillo 1968; French & Paolillo 1976a, b). One could 

assume that it provides protection from UV light and other 

environmental influences, as well as changing the internal 

environment, and that these influences are important in 

shaping the further development of the capsule, as will be 

discussed in another chapter. 

Capsule Development 

In mosses, once the calyptra has been shed, the 

operculum (lid) of the capsule is exposed. This operculum 

must come off before the spores can be dispersed. The 

dehiscence of the operculum is usually facilitated by 

drying of the capsule that causes it to shrink and compress 

the contents. But a few are cleistocarpous (indehiscent; 

lacking a regular means of opening), thus lacking an 

operculum.

Figure 21. Aloina rigida with stalk and capsule and with 

foot imbedded in gametophyte tissue. Photo by Michael Lüth. 

Just under the lid of most moss capsules you will find 

the peristome teeth (in mosses, fringe of teeth around 

opening of capsule, involved in spore dispersal; Figure 22, 

Figure 23). These are usually hygroscopic (responding to 

humidity changes) and may flex back and forth in response 

to moisture changes to aid in gradual dispersal. In most 

cases, these function best as the capsule is drying, but in 

some taxa moisture actually facilitates dispersal. Perhaps 

their best role is in preventing the spores from all exiting 

the capsule at the same time, as happens in the liverworts 

and Sphagnum and most likely also in the mosses with 

valvate capsules. The sporophyte capsule usually has a 

columella (Figure 22) that is columnar like those in 

Polytrichopsida, providing structure. 

Unlike the valvate capsules of liverworts and some 

moss classes, the sporophytes of the Bryopsida are 

photosynthetic. The same pigments often occur in both 

generations: chlorophylls a and b, carotene, lutein, 

violaxanthin, and zeaxanthin (Freeland 1957). Even the 

ratio of chlorophyll a to b is approximately the same – 

about 2.5:1 (Rastorfer 1962). Nevertheless, the 

gametophyte contains a higher chlorophyll concentration 

than does the sporophyte and the ratio of photosynthesis to 

respiration is likewise higher in the gametophyte. Despite 

its photosynthetic abilities, the sporophyte still depends on 

the gametophyte for some of its carbohydrates (Krupa 

1969). 

These stages of the life cycle are summarized in Figure 

24. 


Figure 22. Section of Mnium capsule. This capsule actually 

hangs down, so teeth are on the bottom of the picture. Photo by 

Janice Glime. 

Figure 23. Arthrodontous peristome teeth. Left: peristome of Bartramia pomiformis. Photo by Zen Iwatsuki. Right: spores and 

peristome teeth of Trichostomum showing cell thickenings. Photo by Janice Glime.


Figure 24. Life cycle of the moss Funaria hygrometrica. Drawn by Shelly Meston.

. Chapter 2-7: Bryopsida 53 

Figure 25. Vegetative characters (gametophyte) of Class Bryopsida. Upper Left: Plagiomnium medium stem and leaves. Photo 

by Michael Lüth. Upper right: Plagiomnium stem cross section showing central strand of hydroids. Note smaller darkened areas in 

stem cortex that are leaf traces. Photo by Janice Glime. Middle Left: Leaf of Rhizomnium illustrating a border, small, roundish cells, 

and a distinct costa. Tip of leaf lacking a costa, illustrating elongate cells and undifferentiated apical leaf cells. Photo by Zen Iwatsuki. 

Middle Right: Portion of Plagiomnium leaf showing border. Photo by Janice Glime. Lower Left: Fontinalis stem, leaves, and tuft of 

rhizoids. Photo by Janice Glime. Lower Right: Microscopic view of rhizoids showing single cell thickness and diagonal cross walls. 

Photo by Janice Glime.

54 Chapter 2: The Life Cycle: Surviving Change 

Figure 26. Sexual reproduction of mosses. Upper row shows male reproductive parts. Splash platforms (left) of Mnium hornum 

in which antheridia may be located, or they can be among ordinary leaves (center); among the antheridia are paraphyses (center and 

right) that help in retaining water and in forcing sperm out of the antheridia at maturity. Lower row shows female reproductive parts. 

Perichaetial leaves and young sporophytes of Plagiomnium cuspidatum (left), archegonia from leaf bases of Pleurozium schreberi 

(center), and a section of archegonia (right) with sperm in the neck canal. Plant photos by Michael Lüth; photomicrographs by Janice 

Glime.


Figure 27. Life cycle of a moss such as Mnium (Bryopsida). G represents Gametophyte; S represents Sporophyte. Drawings by 

Allison Slavick, Noris Salazar Allen, and Janice Glime.

56 Chapter 2: The Life Cycle: Surviving Change 

Summary 

The Bryopsida is the largest and most diverse class 

of Bryophyta. In Bryopsida, as in Polytrichopsida, an 

operculum usually covers peristome teeth that often 

aid dispersal. Bryopsida have arthrodontous 

peristome teeth, separating them from the 

Polytrichopsida, which have nematodontous teeth. 

All other classes of Bryobiotina lack peristomes. 

The life cycle of Bryopsida involves a protonema 

that is usually threadlike and develops from the 

germinating spore, developing numerous buds and 

gametophores. Gametophores produce archegonia 

and/or antheridia and the embryo develops within the 

archegonium. 

Sporophytes remain attached to the gametophyte 

and produce spores by meiosis. As in all Bryophyta, 

Bryopsida produce spores from the sporophyte only 

once. 

Vegetative reproduction is common among 

bryophytes. Bryophyta can reproduce by fragments as 

well as specialized asexual structures and thus add a 

new dimension to life cycle strategies. 

Acknowledgments 

I appreciate the comments and suggestions of Karla 

Werner, who offered a beginner's perspective. Noris 

Salazar Allen offered constructive criticisms on the 

taxonomic descriptions and helped with the proof reading 

and life cycle diagrams. 

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