Asterella

Liverwort

General: Asterella is a liverwort, a type of simple, nonvascular plant, akin to mosses. Like mosses, liverworts are
restricted to moist habitats or moist times of the year.

CSUCI: This attractive little liverwort has not yet identified to species. Two species with the appearance are native to
Southern California. One is dioecious (having separate male and female plants); the other is monoecious (having both
male and female function in every plant). Plants at CSUCI have not yet been observed undergoing sexual reproduction,
thus the species diagnosis remains uncertain.

Astrella is found widely around CSUCI, but only in well-shaded habitats (such as the shadiest parts of north facing
slopes). It is active (green) only in the days or weeks immediately following rainfall. When dry and inactive, it is black
and crusty.
BFNA Title: Asterella
Author: M. L. Hicks
Date: March 20, 2003
Edit Level: R Brum+
Version: 1

Bryophyte Flora of North America, Provisional Publication

Missouri Botanical Garden
PO Box 299
St. Louis, MO 63166-0299 USA
www.www.mobot.org/plantscience/BFNA/BFNAmenu.htm

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Asterella - Aytoniaceae

XX. ASTERELLA Palisot de Beauvois, Lam. Dist. Sci. Nat. 3: 257. 1805 • [aster + -ella, little
star, for shape of female receptacle]

Marie L. Hicks

Plants in depressed interwoven patches or partial rosettes, simple or forked; branching

primarily dichotomous or with intercalary branches, arising ventrally by narrow stipes,
clavate, cordate or elongate; apical innovations occasional, forming cordate or elongate
segments. Thalli 20--30 × 1.5--6 mm, margins plane or ascending, often undulate, not
crenulate; dorsal side green with purplish margins, underside purple; drying plant margins
curling upward and inward exposing the purple underside, appearing as blackish-purple
tubes; dorsal epidermis not to very slightly areolate, firm, persistent; cell walls thin to
moderately thick-walled, trigones small or absent; oil cells scattered; air pores simple, the
openings surrounded by 1--3 tiers of 4--8 scarcely differentiated cells; mid-ventral thallus
8--15 cells thick, of compact parenchymatous layers forming a broad keel gradually
diminishing in thickness to the margins; dorsal spongy layers 2--4 cells thick, containing air
chambers lacking photosynthetic filaments; oil cells scattered; rhizoids dense, some
smooth, some with internal pegs; ventral scales purple, in 2 rows, lunate to ovate with
scattered oil cells; appendages hyaline, white or purplish, 1--2(--3) per scale, triangular to
acuminate, not or scarcely constricted at base. Specialized asexual reproduction
absent. Sexual condition monoecious or dioecious; androecia sessile, often purplish, in
dorsal patches of papillae or in discrete thickened pads surrounded by small subulate
scales; the position of androecia in some monoecious species may vary from posterior to
the gynoecial stalks (paroecious) to apical on dichotomous branches or short intercalary
branches (autoecious) or antheridial papillae occasionally form a mid-dorsal streak;
gynoecia terminal on thalli; stalks 0.4--5 cm with one rhizoid furrow, with or without
hairlike scales at bases and/or apices; carpocephala bell-shaped, domed, ovate or
umbonate with air pores on upper surface, developing 1--4 lobes below, each containing
one sporophyte surrounded by an exerted, white to purplish, pleated conical
pseudoperianth that splits into 8--16 apically connate linear segments, becoming free with
age in some species. Sporophyte capsule spherical, irregularly circumscissile; spores
areolate, the areolae bordered by partitions appearing in silhouette as a wing; elaters
120--250 × 12--15 µm, 1--3 spiral.

Species ca. 80 (8 in the flora): on soil worldwide.

SELECTED REFERENCES Evans, A. M. 1920. The North American Species of Asterella.

Contrib. U.S. Nat. Herbarium. 20(8): 247--312. Schuster, R.M. 1992. The Hepaticae and
Anthocerotae of North America. Vol. 6. Chicago.

1. Gynoecial stalks naked, without hairlike scales at bases or apices; plants paroecious or
autoecious .
2. Spores dark brown; southwestern ... 1. Asterella palmeri
2. Spores yellow or pale brown; eastern or boreal.
3. Androecia distinct, paroecious or autoecious; pseudoperianth segments
free at maturity; boreal-alpine .....2. Asterella gracilis
3. Androecia inconspicuous, of few papillae at base of gynoecial stalk;
pseudoperianths conical, segments not free; east of 100th meridian .....3.
Asterella tenella
1. Gynoecial stalks with hairlike scales at bases or apices (or both).
4. Thallus branches primarily lateral intercalary; androecia on short stipitate
branches, occasionally on main thallus; southwestern.
5. Carpocephala upper surfaces with fingerlike protruberances;
stalks with hairlike scales at bases and apices; Texas ......4.
Asterella echinella
5. Carpocephala upper surfaces smooth or with low tubercules;
stalks with hairlike scales at apices, none at bases; west coast ... 5.
Asterella bolanderi
4. Thallus branches primarily dichotomous; androecia on main thalli or its
dichotomous branches.
6. Carpocephala pale green with spreading lobes and long pale
hairlike scales beneath, base of stalk naked; plants dioecious; west
coast .. 6. Asterella californica
6. Carpocephala green or purplish, the lobes directed downward;
stalks with hairlike scales at base; boreal.
7. Spores and elaters purple; stalks with hairlike scales
beneath carpocephala and at base ..7. Asterella
lindenbergiana
 7. Spores yellow or pale brown; without hairlike scales
beneath carpocephala; conspicuous white scale
appendages forming cluster at apices of thallus branches ...
8. Asterella saccata

1. Asterella palmeri (Austin) Underwood, Bot. Gaz. 20: 63. 1895

Fimbriaria palmeri Austin, Bull. Torrey Bot. Cl. 6: 47. 1875

Plants green with purplish undulate margins and deep purple ventral side; branching
dichotomous or thalli simple; intercalary lateral branches rare. Thalli 5--10 × 2--4 mm;
epidermis smooth, cells mostly 30 x 25 µm, thin-walled, with very small trigones, oil cells
absent; air pores surrounded by 1--2 tiers of 5--6 cells with slightly thickened radial walls;
ventral scales imbricate, deep purple with scattered oil cells; appendages 1--2, purplish or
hyaline, subulate to acuminate, often with few small marginal teeth, extending slightly
beyond thallus margins at apices. Sexual condition paroecious, occasionally autoecious;
androecia small dorsal group of papillae at posterior base of gynoecial stalks or
occasionally forming mid-dorsal streaks of papillae; gynoecia terminal on principal thalli;
stalks naked, without hairlike scales at bases or apices, 1--2 cm, purplish; carpocephala
high-domed to ovoid or conical, about 4 mm high, 2.5 mm wide, with 3--4 lobes directed
downward; pseudoperianths white, conelike, cleft 1/3--1/2 the length into 8--10 or more
segments with attached apices. Sporophyte capsules dark brown; spores dark brown,
60--80 µm; elaters curved, brown, 2-spiral, 150--180 µm.

Capsules mature Feb.--Apr. Soil banks in rather dry situations; sw Calif.; Mexico (Baja
California Norte).

This species can be recognized when fruiting by the dark brown spores. Other species
have yellow or purple spores except A. echinella which has pale brown spores and is
unlikely to be confused with A. palmeri because of the strongly tuberculate carpocephala
of the former.

2. Asterella gracilis (F. Weber) Underwood, Bot Gaz. 20: 61. 1895

Marchantia gracilis F. Weber, Hist. Musc. Hep. Prodr. 105. 1815

Plants green with deep purplish undersides and margins; branching dichotomous, ventral
intercalary branches occasional. Thalli 5--15 × 1--3 mm; epidermis smooth, cells
thin-walled, 25--35 × 25--28 µm, trigones small or absent, oil cells few, scattered; air pores
surrounded by 2--3 tiers of 6--8 scarcely differentiated cells; ventral scales purplish with
few oil cells; appendages 1--2, lanceolate. Sexual condition paroecious; androecia
purplish groups of papillae posterior to bases of gynoecial stalks, occasionally on separate
branches, then ovate; gynoecia terminal on dichotomous branches; stalks 1--3 cm, naked
above, bases purplish with inconspicuous, subulate, deciduous scales; carpocephala
hemispheric, 2--3 mm across, upper surface smooth, becoming low tuberculate on drying,
2--4 lobed below; pseudoperianths short, directed obliquely downward, the segments
usually 8, free with age. Sporophyte capsules yellowish-brown; spores yellow, 60--85
µm; elaters yellowish, 2--3 spiral, 150--200 µm.

Capsules mature Apr.--Aug. Moist soil over rock, usually calcareous in Arctic-alpine areas;
Greenland, Alta., B.C., N.W.T., Nun. (Baffin Island); Alaska, Ariz., Calif., Colo., Mich., Minn.,
Mont., N.Mex., Utah, Wyo; Europe; Asia.

This plant has been erroneously known in North America as A. Ludwigii (Schwaegrichen)
Underwood and was listed as such in past publications. According to R. Grolle (1975) the
type of A. ludwigii is Mannia triandra.

3. Asterella tenella (Linnaeus) Palisot de Beauvois, Lam. Dict. Sci. Nat. 3:257. 1805
Marchantia tenella Linnaeus, Sp. Pl. 1137. 1753

Plants green with purplish undulate margins and purplish underside; branching
dichotomous, rarely with intercalary branches. Thalli 6--15 × 1.5--3 mm; epidermis
smooth, cells 25--40 × 20--25 µm, walls not to slightly thickened, trigones absent or small;
oil cells few, scattered; air pores surrounded by 1--3 tiers of 4--6 scarcely differentiated
cells; ventral scales purple with few scattered oil cells; appendages 1--2, white, lanceolate.
Sexual condition paroecious, androecia of inconspicuous papillae at posterior base of
gynoecial stalk; gynoecia in apical notch of main thalli; stalks naked, often purplish, 1--3
cm, without hairlike scales at bases or apices; carpocephala bell shaped, smooth, 3--4 mm
across, lobes 3--4, short, directed downward; pseudoperianths white or purplish, conical, of
8--12 segments, connate at their apices, occasionally free with age. Sporophyte
capsules pale yellowish brown; spores yellow, 85--100 µm; elaters yellowish-brown,
2-spiral, 150--250 µm.

Capsules mature Mar.--June. Soil over rock near streams; low elevations; Ont.; Que., Ala.,
Ark., Conn., D.C., Ga., Ill., Ind., Kans., Ky., La., Maine, Md., Mass., Miss., Mo., N.H., N.J., N.C.,
N.Y., Ohio, Okla., Pa., R.I., S.C., Tenn., Tex., Vt., Va., W.Va.

This species is common in e North America extending westward to Kansas; it has not been
found west of the 100th meridian.

4. Asterella echinella (Gottsche) Underwood, Bot. Gaz. 20: 62. 1895

Fimbriaria echinella Gottsche, Dansk. Vid. Sellsk. Skrift. V 6: 367 1863

Plants green above with deep purple pigmentation along margins and below; branching
intercalary from a simple or dichotomous main thallus; lateral-ventral branches frequent,
arising by narrow stalks, the distal portions often obcordate; apical innovations also occur.
Thalli 10--30 × 2--3 mm with margins often undulating and ascending; epidermis smooth,
cells 30--50 × 30--35 µm with slightly thickened walls, without distinct trigones; air pores
surrounded by 1--2 tiers of 6--8 cells; ventral scales deep purple with scattered oil cells;
appendages 1--2, white or purplish tinged, subulate to acuminate, 3--4 cells wide, slightly
narrowed at base. Sexual condition autoecious, male branches short, obcordate,
sometimes innovating apically; androecia terminal, deep purple, oblong, ovate or
subcordate, thickened, the margins with small subulate scales; gynoecia on principal
branches or short innovative branches; stalks 0.5--1.5 cm, reddish tinged with hairlike
scales at apices and sparse scales at bases; carpocephala domed, 2--3 mm wide with
conspicuous finger-like tubercules on upper surfaces, 1--3 lobed below; pseudoperianths
white to slightly purple, of 8--10 segments, connate at apices. Sporophyte capsules
brownish; spores yellowish to pale brown, 60--100 µm, elaters brown, 1--2 spiral, 140--200
µm.

Capsules mature May. Soil over rocks, often calcareous, near streams; Tex.; Mexico.

The conspicuous finger-like tubercules on carpocephala distinguish this species from

others in the flora. Asterella echinella was listed by W. S. Sullivant (1856) and L. M.
Underwood (1884) as A. elegans, a plant of Mexico. Subsequently R. Stotler and B.
Crandall-Stotler (1977) listed it as A. elegans subsp. echinella (Gottsche) Del Rosario in
error. Thus far, A. elegans has not been found north of Mexico.

5. Asterella bolanderi (Austin) Underwood, Bot. Gaz. 20: 61 1895

Fimbriaria bolanderi Austin, Proc. Acad. Sci. Phila. 21: 230 1869

Plants green above, deep purple below and along margins, which are undulate and
slightly ascending; branching by numerous short intercalary lateral branches that arise
beneath main thallus; dichotomous branches few. Thalli 10--30 × 2--4 mm; epidermis
smooth, cells 30--40 × 20--30 µm, the walls slightly thickened, trigones small or lacking, oil
cells few, scattered; air pores scarcely elevated, with 2--3 tiers of 7--8 cells around the
opening; ventral scales deep purplish-red with scattered oil cells; appendages 1--2 purplish
or hyaline, lanceolate to acuminate. Sexual condition autoecious, sex organs on short
latero-ventral intercalary branches; androecia on very short clavate branches, the
androecia ovate, without surrounding scales; gynoecia on short, obcordate branches,
stalks reddish, 1--3 cm with sparse hair-like scales at apices, few or none at bases;
carpocephala high domed and bell-shaped, 2.4--4 mm across, usually 4-lobed, the lobes
directed obliquely downward; pseudoperianths white or reddish tinged, 10--12 cleft, the
segments connate at apices. Sporophyte capsules yellowish brown; spores yellow to
pale brown, 65--100 µm; elaters yellow to pale brownish, 2(--3)-spiral, 150--220 µm.

Capsules mature Mar.--May. Soil of shaded banks; Calif., sw Oreg.

The presence of frequent, short, intercalary branches and paucity of dichotomous

branching distinguishes this species from others of the genus in the flora. The principal
thalli often curl when dry, exposing the dark purple underside. Asterella bolanderi subsp.
acrogyna R. M. Schuster named from a single Texas collection (R. M. Schuster 1985), was
a specimen of Reboulia, later treated (R. M. Schuster 1992) as Reboulia hemisphaerica
subsp. acrogyna (R. M. Schuster) R. M. Schuster.

6. Asterella californica (Underwood) Underwood, Bot. Gaz, 20: 60 1895

Fimbriaria californica Underwood, Bull. Ill. Lab. Nat. Hist. 2: 41 1884

Plants pale green to green dorsally with purple ascending margins and dark purple
undersides, edges tending to curl upward exposing the dark underside when dry;
branching dichotomous, seldom with intercalary branches. Thalli 10--25 × 4--10 mm;
epidermis faintly areolate, cells 50 × 30 µm with thin walls, trigones small or lacking, oil
cells few, scattered; air pores surrounded by 2--3 tiers of 6--7 cells with slightly thickened
radial walls; ventral scales purplish with few oil cells; appendages 1--3, hyaline,
triangular-acuminate. Sexual condition dioecious; separate male plants often
intermingled with female plants; androecia dorsal, forming thick, distinct ovate or elongate
patches, sometimes with subulate scales around the margins; gynoecia terminal on thalli,
stalks slightly purplish, 1--3 cm with pale, long, fine, hairlike scales at apices, none at
bases; carpocephala pale green, 4--5 mm across, hemispheric, becoming umbonate with
age, distinctly lobed with 3--4(--5) lobes directed horizontally outward; pseudoperianths
conical, white with 12 or more segments, connate at apices. Sporophyte capsules
yellowish; spores yellow, 100--120 µm; elaters yellowish, 1--2 spiral, 250--300 µm.

Capsules mature Jan.--May. Soil of shaded banks in rather dry areas; Ariz. (Gila Co.), Calif.,
sw Oreg.; Mexico (Baja California Norte).

This is the only dioecious species in the flora. The female receptacles are large for the size
of the plant and the pseudoperianths are directed outward horizontally rather than
downward. The pale color of the carpocephala is in sharp contrast with the inrolled thalli
covering the green upper surface exposing the dark purple underside.

7. Asterella lindenbergiana (Corda) Lindberg, Musc. Scand. 1. 1879

Fimbriaria lindenbergiana A. Corda in C. Nees von Esenbeck, Naturg. Eur. Leberm. 4: 283.
1838

Plants green, often pigmented with reddish-purple blotches above, purplish-red along the
undulate, ascending margins and below; branching dichotomous, ventral intercalary
branches infrequent. Thalli 10--30 × 4--6 mm; epidermis faintly areolate, cells 30 × 25
µm with thin walls and no trigones, oil cells few, scattered; air pores surrounded by 3--4
tiers of 6--8 scarcely differentiated cells; ventral scales large, purple with scattered oil
cells; appendages 1--2, white, lanceolate to acuminate. Sexual condition paroecious or
autoecious, androecia posterior to gynoecial stalks or on separate branches, ovate, with
variable number of papillae, or forming dorsal streaks of papillae; gynoecia terminal on
main thalli; stalks purplish, 1.5--2.5 cm with hairlike scales at bases and apices;
carpocephala 3--4 mm across, conical with low tubercules above and 3--4 short lobes
directed downward; pseudoperianths pleated, often purplish, of 12 or more segments with
connate apices. Sporophyte capsules purple; spores deep purple, 80--100 µm; elaters
purple, 2-spiral, 100--150 µm.

Capsules mature July--Aug. Damp, mossy soil, frequently calcareous, arctic-alpine; Alta.,
B.C.; Alaska, Colo., Mont., Oreg., Utah, Wash., Wyo.; Mexico; South America (Colombia);
Europe.

The purple spores and elaters of this species are distinctive and when fruiting it can be
readily separated from other species in the flora on that basis.

8. Asterella saccata (Wahlenberg) A. Evans, Contr. U.S. Nat Herb. 20: 276 1920

Marchantia saccata Wahlenberg, Ges. Nat. Freund. Berlin Mag. 5: 296 1811

Plants green above, purplish along margins and underneath, the thalli curl upward around
apical margins exposing white scale appendages, forming a conspicuous apical tuft;
branching dichotomous, ventral intercalary branches infrequent. Thalli 5--10 × 2--3 mm;
epidermis smooth, cells 20 × 30 µm with slightly thickened walls and small trigones; air
pores indistinct, surrounded by 1--3 tiers of 5--7 scarcely differentiated cells, oil cells few,
scattered; ventral scales deep purple, long tapered, with scattered oil cells; appendages
1--2, white, long tapered lanceolate to acuminate, as long or longer than scales, curling up
around the anterior thalli margins, forming conspicuous white clusters at apices. Sexual
condition paroecious, occasionally autoecious; androecia form ill-defined dorsal streaks of
papillae at posterior bases of gynoecial stalks or on nearby branches; gynoecia terminal on
main thalli, stalks 1--2 cm, purplish with cluster of white hairlike scales at bases and none
at apices; carpocephala 2--3 mm across, ovate, at least 1-1/2 times taller than wide with
3--4 lobes directed downward; pseudoperianths conical, of 6--8 white segments, connate
at apices. Sporophyte capsules yellowish; spores yellow to pale brown, 80--90 µm;
elaters yellowish, 1--3 spiral, 150--200 µm.

Capsules mature May. Soil in rock crevices, usually calcareous in arctic-alpine areas;
Greenland; Alta.; Alaska, Minn., Mont., Wash.; Europe; Asia.

The white scale appendages exposed at thalli apices and along upturned margins are
distinctive. These scale clusters have caused confusion with Mannia fragans, which has a
similar apical cluster. Unconfirmed reports from areas south of this species' range may
represent M. fragans, which also lacks a pseudoperianth and has brown, not yellow spores.

OTHER REFERENCES

Grolle, R. 1975. Miscellanea hepaticologica (141-150) J. Bryol. 8: 483--492.

Schuster, R. M. 1985. Some new taxa of Hepaticae. Phytologia 57: 410.

Schuster, R. M. 1992. The Hepaticae and Anthocerotae of North America. Chicago.

Stotler, R. and B. Crandall-Stotler. 1977. A checklist of the liverworts and hornworts of

North America. Bryologist 80: 405--428.

Sullivant, W. S. 1856. The musci and hepaticae of the United States east of the Mississippi
River. In: A. Gray, Gray's Manual of Botany Ed. 2, pp. 607--737, pls 1--8.

Underwood, L. M. 1884. Descriptive Catalogue of the North American hepaticae, North of

Mexico. Bull. Illinois State Lab. Nat. Hist. 2: 1-133.
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BRYOPHYTES

Plant scientists recognize two kinds of land plants, namely, bryophytes, or nonvascular land
plants and tracheophytes,or vascular land plants. Bryophytes are small, herbaceous plants
that grow closely packed together in mats or cushions on rocks, soil, or as epiphytes on the
trunks and leaves of forest trees. Bryophytes are distinguished from tracheophytes by two
important characters. First, in all bryophytes the ecologically persistent, photosynthetic phase
of the life cycle is the haploid, gametophyte generation rather than the diploid sporophyte;
bryophyte sporophytes are very short-lived, are attached to and nutritionally dependent on
their gametophytes and consist of only an unbranched stalk, or seta, and a single, terminal
sporangium. Second, bryophytes never form xylem tissue, the special lignin- containing, water-
conducting tissue that is found in the sporophytes of all vascular plants. At one time,
bryophytes were placed in a single phylum, intermediate in position between algae and
vascular plants. Modern studies of cell ultrastructure and molecular biology, however,confirm
that bryophytes comprise three separate evolutionary lineages, which are today recognized as
mosses (phylum Bryophyta), liverworts (phylum Marchantiophyta) and hornworts (phylum
Anthocerotophyta). Following a detailed analysis of land plant relationships, Kenrick and Crane
(1998) proposed that the three groups of bryophytes represent a grade or structural level in
plant evolution, identified by their "monosporangiate" life cycle. Within this the geologically
oldest group, sharing a fossil record with the oldest vascular plants in the Devonian era.

Of the three phyla of bryophytes, greatest species diversity is found in the mosses, with up to
15,000 species recognized. A moss begins its life cycle when haploid spores, which are
produced in the sporophyte capsule, land on a moist substrate and begin to germinate. From
the one-celled spore, a highly branched system of filaments, called the protonema, develops.
Cell specialization occurs within the protonema to form a horizontal system of reddish-brown,
anchoring filaments, called caulonemal filaments and upright, green filaments, called
chloronemal filaments. Each protonema, which superficially resembles a filamentous alga, can
spread over several centimeters to form a fuzzy green film over its substrate. As the
protonema grows, some cells of the caulonemal filaments specialize to form leafy buds that
will ultimately form the adult gametophyte shoots. Numerous shoots typically develop from
each protonema so that, in fact, a single spore can give rise to a whole clump of moss plants.
Each leafy shoot continues to grow apically, producing leaves in spiral arrangement on an
elongating stem. In many mosses the stem is differentiated into a central strand of thin-walled
water-conducting cells, called hydroids, surrounded by a parenchymatous cortex and a thick-
walled epidermis. The leaves taper from a broad base to a pointed apex and have lamina that
are only one-cell layer thick. A hydroid-containing midvein often extends from the stem into
the leaf. Near the base of the shoot, reddish-brown, multicellular rhizoids emerge from the
stem to anchor the moss to its substrate. Water and mineral nutrients required for the moss to
grow are absorbed, not by the rhizoids,but rather by the thin leaves of the plant as rain water
washes through the moss cushion.

As is typical of bryophytes, mosses produce large, multicellular sex organs for reproduction.
Many bryophytes are unisexual, or sexually dioicous. In mosses male sex organs, called
antheridia, are produced in clusters at the tips of shoots or branches on the male plants and
female sex organs, the archegonia, are produced in similar fashion on female plants.
Numerous motile sperm are produced by mitosis inside the brightly colored, club-shaped
antheridia while a single egg develops in the base of each vase-shaped archegonium. As the
sperm mature, the antheridium swells and bursts open. Drops of rain water falling into the
cluster of open antheridia splash the sperm to near-by females. Beating their two whiplash
flagellae, the sperm are able to move short distances in the water film that covers the plants
to the open necks of the archegonia. Slimey mucilage secretions in the archegonial neck help
pull the sperm downward to the egg. The closely packed arrangement of the individual moss
plants greatly facilitates fertilization. Rain forest bryophytes that hang in long festoons from
the trees rely on torrential winds with the rain to transport their sperm from tree to tree, while
the small pygmy mosses of exposed, ephemeral habitats depend on the drops of morning dew
to move their sperm.Regardless of where they grow, all bryophytes require water for sperm
dispersal and subsequent fertilization.

Embryonic growth of the sporophyte begins within the archegonium soon after fertilization. At
its base, or foot, the growing embryo forms a nutrient transfer zone, or placenta, with the
gametophyte. Both organic nutrients and water move from the gametophyte into the
sporophyte as it continues to grow. In mosses the sporophyte stalk, or seta, tears the
archegonial enclosure early in development, leaving only the foot and the very base of the
seta embedded in the gametophyte. The upper part of the archegonium remains over the tip
of the sporophyte as a cap-like calyptra. Sporophyte growth ends with the formation of a
sporangium or capsule at the tip of the seta. Within the capsule, water-resistant spores are
formed by meiosis. As the mature capsule swells, the calyptra falls away. This allows the
capsule to dry and break open at its tip. Special membranous structures, called peristome
teeth, that are folded down into the spore mass,now bend outward, flinging the spores into the
drying winds. Moss spores can travel great distances on the winds, even moving between
continents on the jet streams. Their walls are highly protective, allowing some spores to
remain viable for up to 40 years. Of course, if the spore lands in a suitable, moist habitat,
germination will begin the cycle all over again.

Liverworts and hornworts are like mosses in the fundamental features of their life cycle, but
differ greatly in organization of their mature gametophytes and sporophytes. Liverwort
gametophytes can be either leafy shoots or flattened thalli. In the leafy forms, the leaves are
arranged on the stem in one ventral and two lateral rows or ranks, rather than in spirals like
the mosses. The leaves are one cell layer thick throughout, never have a midvein and are
usually divided into two or more parts called lobes. The ventral leaves, which actually lie
against the substrate, are usually much smaller than the lateral leaves and are hidden by the
stem. Anchoring rhizoids, which arise near the ventral leaves, are colorless and unicellular. The
flattened ribbon-like to leaf-like thallus of the thallose liverworts can be either simple or
structurally differentiated into a system of dorsal air chambers and ventral storage tissues. In
the latter type, the dorsal epidermis of the thallus is punctuated with scattered pores that
open into the air chambers. Liverworts synthesize a vast array of volatile oils, which they store
in unique organelles called oil bodies. These compounds impart an often spicy aroma to the
plants and seem to discourage animals from feeding on them. Many of these compounds have
potential as antimicrobial or anticancer pharmecuticals.

Liverwort sporophytes develop completely enclosed within gametophyte tissues until their
capsules are ready to open. The seta, which is initially very short,consists of small, thin-walled,
hyaline cells. Just prior to capsule opening, the seta cells lengthen, thereby increasing the
length of the seta upto 20 times its original dimensions. This rapid elongation pushes the
darkly pigmented capsule and upper part of the whitish seta out of the gametophytic tissues.
With drying, the capsule opens by splitting into four segments, or valves. The spores are
dispersed into the winds by the twisting motions of numerous intermixed sterile cells, called
elaters. In contrast to mosses, which disperse their spores over several days, liverworts
disperse the entire spore mass of a single capsule in just a few minutes.
Hornworts resemble some liverworts in having simple, unspecialized thalloid gametophytes,
but they differ in many other characters. For example, colonies of the symbiotic
cyanobacterium Nostoc fill small cavities that are scattered throughout the ventral part of the
hornwort thallus. When the thallus is viewed from above, these colonies appear as scattered
blue-green dots. The cyanobacterium converts nitrogen gas from the air into ammonium,
which the hornwort requires in its metabolism and the hornwort secretes carbohydrate-
containing mucilage which supports the growth of the cyanobacterium.Hornworts also differ
from all other land plants in having only one large, algal-like chloroplast in each thallus cell.
Hornworts get their name from their long, horn-shaped sporophytes. As in other bryophytes,
the sporophyte is anchored in the gametophyte by a foot through which nutrient transfer from
gametophyte to sporophyte occurs. The rest of the sporophyte, however, is actually an
elongate sporangium in which meiosis and spore development take place. At the base of the
sporangium, just above the foot, is a mitotically active meristem,which adds new cells to the
spore-producing zone throughout the life span of the sporophyte. In fact, the sporangium can
be releasing spores at its apex, at the same time that new spores are being produced by
meiosis at its base. Spore release in hornworts takes place gradually over a long period of
time, and the spores are mostly dispersed by water movements rather than by wind

Mosses, liverworts and hornworts are found throughout the world in a variety of habitats. They
flourish particularly well in moist, humid forests like the fog forests of the Pacific northwest or
the montane rain forests of the southern hemisphere. Their ecological roles are many.They
provide seed beds for the larger plants of the community, they capture and recycle nutrients
that are washed with rainwater from the canopy and they bind the soil to keep it from eroding.
In the northern hemisphere peatlands, wetlands often dominated by the moss Sphagnum, are
particularly important bryophyte communities. This moss has exceptional water-holding
capacity, and when dried and compressed, forms a coal-like fuel. Throughout northern Europe,
Asia and North America, peat has been harvested for centuries for both fuel consumption and
horticultural uses and today peatlands are managed as a sustainable resource.

BIBLIOGRAPHY

• Crandall-Stotler, Barbara. "Morphogenetic designs and a theory of

bryophyte origins and divergence." BioScience 30(1980): 580-585.

• Hébant, Charles. The Conducting Tissues of Bryophytes. Vaduz: J.

Cramer, 1977.

• Kenrick, Paul, and Peter R. Crane. The Origin and Early Diversification
of Land Plants: A Cladistic Study. Washington, D. C.: Smithsonian Institution
Press, 1997.

• Miller, Norton G. "Bogs, bales and BTUs: a primer on peat." Horticulture

59 (1981): 38-45.

• Schofield, W. B. Introduction to Bryology. New York: Macmillan, 1985.

• Shaw, Jonathon A., and Bernard Goffinet, eds. The Biology of

Bryophytes. Cambridge, England: Cambridge University Press, 2000

Distinguishing Characters of Mosses (Phylum Bryophyta), Liverworts (Phylum

Marchantiophyta) and Hornworts
(Phylum Anthocerotophyta)
Character Bryophyta Marchantiophyta Anthocerotophyta

Filamentous, forming Globose, forming one Globose, forming one

Protonema
many buds bud bud

Leafy shoot or thallus;

Gametophyte form Leafy shoot thallus simple or with Simple thallus
air chambers

Leaf arrangement Leaves in spirals Leaves in three rows Not Applicable

Leaves undivided, Leaves divided into 2+

Leaf form Not Applicable
midvein present lobes, no midvein

Single plastids with

Special organelles None Oil bodies
pyrenoids

Present in both
Present only in a few
Water conducting cells gametophytes and Absent
simple thalloid forms
sporophytes

Rhizoids Brown, multicellular Hyaline, one-celled Hyaline, one-celled

Apical clusters (leafy

Sunken in thallus,
Gametangial position Apical clusters forms) or on upper
scattered
surface of thallus

Present in both
Present on sporophyte Absent in both
Stomates sporophyte and
capsule generations
gametophyte

Photosynthetic,
emergent from Hyaline, elongating just
Seta Absent
gametophyte early in prior to spore release
development

Undifferentiated, horn-
Complex with Undifferentiated,
shaped; growing
Capsule operculum, theca and spherical or elongate; of
continuously from a
neck; of fixed size fixed size
basal meristem

Spirally thickened Columella and

Sterile cells in capsule Columella
elaters pseudoelaters

At operculum and
Capsule dehiscence Into 4 valves Into 2 valves
peristome teeth
Prepared by : Barbara Crandall-Stotler, Department of Plant Biology, Southern
Illinois University,
Carbondale, IL 62901-6509

Created by Dr. Raymond E. Stotler and Dr. Barbara J. Crandall-Stotler,

Department of Plant Biology

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BRYOPHYTES

Plant scientists recognize two kinds of land plants, namely, bryophytes, or nonvascular land
plants and tracheophytes,or vascular land plants. Bryophytes are small, herbaceous plants
that grow closely packed together in mats or cushions on rocks, soil, or as epiphytes on the
trunks and leaves of forest trees. Bryophytes are distinguished from tracheophytes by two
important characters. First, in all bryophytes the ecologically persistent, photosynthetic phase
of the life cycle is the haploid, gametophyte generation rather than the diploid sporophyte;
bryophyte sporophytes are very short-lived, are attached to and nutritionally dependent on
their gametophytes and consist of only an unbranched stalk, or seta, and a single, terminal
sporangium. Second, bryophytes never form xylem tissue, the special lignin- containing, water-
conducting tissue that is found in the sporophytes of all vascular plants. At one time,
bryophytes were placed in a single phylum, intermediate in position between algae and
vascular plants. Modern studies of cell ultrastructure and molecular biology, however,confirm
that bryophytes comprise three separate evolutionary lineages, which are today recognized as
mosses (phylum Bryophyta), liverworts (phylum Marchantiophyta) and hornworts (phylum
Anthocerotophyta). Following a detailed analysis of land plant relationships, Kenrick and Crane
(1998) proposed that the three groups of bryophytes represent a grade or structural level in
plant evolution, identified by their "monosporangiate" life cycle. Within this the geologically
oldest group, sharing a fossil record with the oldest vascular plants in the Devonian era.

Of the three phyla of bryophytes, greatest species diversity is found in the mosses, with up to
15,000 species recognized. A moss begins its life cycle when haploid spores, which are
produced in the sporophyte capsule, land on a moist substrate and begin to germinate. From
the one-celled spore, a highly branched system of filaments, called the protonema, develops.
Cell specialization occurs within the protonema to form a horizontal system of reddish-brown,
anchoring filaments, called caulonemal filaments and upright, green filaments, called
chloronemal filaments. Each protonema, which superficially resembles a filamentous alga, can
spread over several centimeters to form a fuzzy green film over its substrate. As the
protonema grows, some cells of the caulonemal filaments specialize to form leafy buds that
will ultimately form the adult gametophyte shoots. Numerous shoots typically develop from
each protonema so that, in fact, a single spore can give rise to a whole clump of moss plants.
Each leafy shoot continues to grow apically, producing leaves in spiral arrangement on an
elongating stem. In many mosses the stem is differentiated into a central strand of thin-walled
water-conducting cells, called hydroids, surrounded by a parenchymatous cortex and a thick-
walled epidermis. The leaves taper from a broad base to a pointed apex and have lamina that
are only one-cell layer thick. A hydroid-containing midvein often extends from the stem into
the leaf. Near the base of the shoot, reddish-brown, multicellular rhizoids emerge from the
stem to anchor the moss to its substrate. Water and mineral nutrients required for the moss to
grow are absorbed, not by the rhizoids,but rather by the thin leaves of the plant as rain water
washes through the moss cushion.

As is typical of bryophytes, mosses produce large, multicellular sex organs for reproduction.
Many bryophytes are unisexual, or sexually dioicous. In mosses male sex organs, called
antheridia, are produced in clusters at the tips of shoots or branches on the male plants and
female sex organs, the archegonia, are produced in similar fashion on female plants.
Numerous motile sperm are produced by mitosis inside the brightly colored, club-shaped
antheridia while a single egg develops in the base of each vase-shaped archegonium. As the
sperm mature, the antheridium swells and bursts open. Drops of rain water falling into the
cluster of open antheridia splash the sperm to near-by females. Beating their two whiplash
flagellae, the sperm are able to move short distances in the water film that covers the plants
to the open necks of the archegonia. Slimey mucilage secretions in the archegonial neck help
pull the sperm downward to the egg. The closely packed arrangement of the individual moss
plants greatly facilitates fertilization. Rain forest bryophytes that hang in long festoons from
the trees rely on torrential winds with the rain to transport their sperm from tree to tree, while
the small pygmy mosses of exposed, ephemeral habitats depend on the drops of morning dew
to move their sperm.Regardless of where they grow, all bryophytes require water for sperm
dispersal and subsequent fertilization.

Embryonic growth of the sporophyte begins within the archegonium soon after fertilization. At
its base, or foot, the growing embryo forms a nutrient transfer zone, or placenta, with the
gametophyte. Both organic nutrients and water move from the gametophyte into the
sporophyte as it continues to grow. In mosses the sporophyte stalk, or seta, tears the
archegonial enclosure early in development, leaving only the foot and the very base of the
seta embedded in the gametophyte. The upper part of the archegonium remains over the tip
of the sporophyte as a cap-like calyptra. Sporophyte growth ends with the formation of a
sporangium or capsule at the tip of the seta. Within the capsule, water-resistant spores are
formed by meiosis. As the mature capsule swells, the calyptra falls away. This allows the
capsule to dry and break open at its tip. Special membranous structures, called peristome
teeth, that are folded down into the spore mass,now bend outward, flinging the spores into the
drying winds. Moss spores can travel great distances on the winds, even moving between
continents on the jet streams. Their walls are highly protective, allowing some spores to
remain viable for up to 40 years. Of course, if the spore lands in a suitable, moist habitat,
germination will begin the cycle all over again.

Liverworts and hornworts are like mosses in the fundamental features of their life cycle, but
differ greatly in organization of their mature gametophytes and sporophytes. Liverwort
gametophytes can be either leafy shoots or flattened thalli. In the leafy forms, the leaves are
arranged on the stem in one ventral and two lateral rows or ranks, rather than in spirals like
the mosses. The leaves are one cell layer thick throughout, never have a midvein and are
usually divided into two or more parts called lobes. The ventral leaves, which actually lie
against the substrate, are usually much smaller than the lateral leaves and are hidden by the
stem. Anchoring rhizoids, which arise near the ventral leaves, are colorless and unicellular. The
flattened ribbon-like to leaf-like thallus of the thallose liverworts can be either simple or
structurally differentiated into a system of dorsal air chambers and ventral storage tissues. In
the latter type, the dorsal epidermis of the thallus is punctuated with scattered pores that
open into the air chambers. Liverworts synthesize a vast array of volatile oils, which they store
in unique organelles called oil bodies. These compounds impart an often spicy aroma to the
plants and seem to discourage animals from feeding on them. Many of these compounds have
potential as antimicrobial or anticancer pharmecuticals.

Liverwort sporophytes develop completely enclosed within gametophyte tissues until their
capsules are ready to open. The seta, which is initially very short,consists of small, thin-walled,
hyaline cells. Just prior to capsule opening, the seta cells lengthen, thereby increasing the
length of the seta upto 20 times its original dimensions. This rapid elongation pushes the
darkly pigmented capsule and upper part of the whitish seta out of the gametophytic tissues.
With drying, the capsule opens by splitting into four segments, or valves. The spores are
dispersed into the winds by the twisting motions of numerous intermixed sterile cells, called
elaters. In contrast to mosses, which disperse their spores over several days, liverworts
disperse the entire spore mass of a single capsule in just a few minutes.

Hornworts resemble some liverworts in having simple, unspecialized thalloid gametophytes,

but they differ in many other characters. For example, colonies of the symbiotic
cyanobacterium Nostoc fill small cavities that are scattered throughout the ventral part of the
hornwort thallus. When the thallus is viewed from above, these colonies appear as scattered
blue-green dots. The cyanobacterium converts nitrogen gas from the air into ammonium,
which the hornwort requires in its metabolism and the hornwort secretes carbohydrate-
containing mucilage which supports the growth of the cyanobacterium.Hornworts also differ
from all other land plants in having only one large, algal-like chloroplast in each thallus cell.
Hornworts get their name from their long, horn-shaped sporophytes. As in other bryophytes,
the sporophyte is anchored in the gametophyte by a foot through which nutrient transfer from
gametophyte to sporophyte occurs. The rest of the sporophyte, however, is actually an
elongate sporangium in which meiosis and spore development take place. At the base of the
sporangium, just above the foot, is a mitotically active meristem,which adds new cells to the
spore-producing zone throughout the life span of the sporophyte. In fact, the sporangium can
be releasing spores at its apex, at the same time that new spores are being produced by
meiosis at its base. Spore release in hornworts takes place gradually over a long period of
time, and the spores are mostly dispersed by water movements rather than by wind

Mosses, liverworts and hornworts are found throughout the world in a variety of habitats. They
flourish particularly well in moist, humid forests like the fog forests of the Pacific northwest or
the montane rain forests of the southern hemisphere. Their ecological roles are many.They
provide seed beds for the larger plants of the community, they capture and recycle nutrients
that are washed with rainwater from the canopy and they bind the soil to keep it from eroding.
In the northern hemisphere peatlands, wetlands often dominated by the moss Sphagnum, are
particularly important bryophyte communities. This moss has exceptional water-holding
capacity, and when dried and compressed, forms a coal-like fuel. Throughout northern Europe,
Asia and North America, peat has been harvested for centuries for both fuel consumption and
horticultural uses and today peatlands are managed as a sustainable resource.

BIBLIOGRAPHY

• Crandall-Stotler, Barbara. "Morphogenetic designs and a theory of

bryophyte origins and divergence." BioScience 30(1980): 580-585.

• Hébant, Charles. The Conducting Tissues of Bryophytes. Vaduz: J.

Cramer, 1977.
 • Kenrick, Paul, and Peter R. Crane. The Origin and Early Diversification
of Land Plants: A Cladistic Study. Washington, D. C.: Smithsonian Institution
Press, 1997.

• Miller, Norton G. "Bogs, bales and BTUs: a primer on peat." Horticulture

59 (1981): 38-45.

• Schofield, W. B. Introduction to Bryology. New York: Macmillan, 1985.

• Shaw, Jonathon A., and Bernard Goffinet, eds. The Biology of

Bryophytes. Cambridge, England: Cambridge University Press, 2000

Distinguishing Characters of Mosses (Phylum Bryophyta), Liverworts (Phylum

Marchantiophyta) and Hornworts
(Phylum Anthocerotophyta)

Character Bryophyta Marchantiophyta Anthocerotophyta

Filamentous, forming Globose, forming one Globose, forming one

Protonema
many buds bud bud

Leafy shoot or thallus;

Gametophyte form Leafy shoot thallus simple or with Simple thallus
air chambers

Leaf arrangement Leaves in spirals Leaves in three rows Not Applicable

Leaves undivided, Leaves divided into 2+

Leaf form Not Applicable
midvein present lobes, no midvein

Single plastids with

Special organelles None Oil bodies
pyrenoids

Present in both
Present only in a few
Water conducting cells gametophytes and Absent
simple thalloid forms
sporophytes

Rhizoids Brown, multicellular Hyaline, one-celled Hyaline, one-celled

Apical clusters (leafy

Sunken in thallus,
Gametangial position Apical clusters forms) or on upper
scattered
surface of thallus

Present in both
Present on sporophyte Absent in both
Stomates sporophyte and
capsule generations
gametophyte
 Photosynthetic,
emergent from Hyaline, elongating just
Seta Absent
gametophyte early in prior to spore release
development

Undifferentiated, horn-
Complex with Undifferentiated,
shaped; growing
Capsule operculum, theca and spherical or elongate; of
continuously from a
neck; of fixed size fixed size
basal meristem

Spirally thickened Columella and

Sterile cells in capsule Columella
elaters pseudoelaters

At operculum and
Capsule dehiscence Into 4 valves Into 2 valves
peristome teeth

Prepared by : Barbara Crandall-Stotler, Department of Plant Biology, Southern

Illinois University,
Carbondale, IL 62901-6509

Created by Dr. Raymond E. Stotler and Dr. Barbara J. Crandall-Stotler,

Department of Plant Biology

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Title: The Classification of New Zealand Hepaticae

Author: E. Amy Hodgson

In: Tuatara: Volume 3, Issue 1, May 1950

Part of: Tuatara : Journal of the Biological Society

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Tuatara: Volume 3, Issue 1, May 1950

17. Asterella australis × 16, F carpocephalum showing laciniate perianth, × 4. — 18.

Targionia hypophylla with terminal valves, × 1½. — 19. Riccardia marginata
showing absence of nerve and lateral calyptra, × 3. — 20. Anthocercs with
linear dehiscing capsule, nat. size. — 21. Marchantia with male carpocephalum
and rounded gammae cups; G. female carpocephalum showing involucres
between the rays, nat. size. — 22. Metzgeria furcata dorsal with ventral
calyptras showing, also mid-rib and ciliate margins; H. ventral with male
branches, x. 4. — 23. Reboulia hemisphaerica, carpocephalum, underneath view
showing bi-valved involucres, × 2½. — 24. Plagiochosma australe with invol.
lobes opening vertically, × 2½. — 26. Lunularia cruciata with lunate gemmae
cups, nat. size. — 27. Sypmhyogyna hymenophyllum showing involucral scales
and calyptras; I. sterile archegonia adhering to calyptra, × 2. — 28. Pallavicinia
Lyellii, J. invol. cup, K. perianth with calyptra included, nat. size. — 29.
Hymenophytum phyllanthus, L. basal ventral invol. cup, M. perianth, calyptra
included, nat. size. — 30. Dehisecnt capsules; N. Taxilejeunea Colensoana
showing valves not divided to the base, × 10; O. Plagiochila × 7; P.
Fossombronia × 7; Q. Metzgeria furcata with elaterophones × 10; R. Asterella ×
5

Previous Figure | Table of Contents | Figure in Context | Next Figure

17. Asterella australis × 16, F carpocephalum showing laciniate perianth, × 4.
18. Targionia hypophylla with terminal valves, × 1½.
19. Riccardia marginata showing absence of nerve and lateral calyptra, × 3.
20. Anthocercs with linear dehiscing capsule, nat. size.
21. Marchantia with male carpocephalum and rounded gammae cups; G. female
carpocephalum showing involucres between the rays, nat. size.
22. Metzgeria furcata dorsal with ventral calyptras showing, also mid-rib and ciliate margins; H.
ventral with male branches, x. 4.
23. Reboulia hemisphaerica, carpocephalum, underneath view showing bi-valved involucres, ×
2½.
24. Plagiochosma australe with invol. lobes opening vertically, × 2½.
26. Lunularia cruciata with lunate gemmae cups, nat. size.
27. Sypmhyogyna hymenophyllum showing involucral scales and calyptras; I. sterile
archegonia adhering to calyptra, × 2.
28. Pallavicinia Lyellii, J. invol. cup, K. perianth with calyptra included, nat. size.
29. Hymenophytum phyllanthus, L. basal ventral invol. cup, M. perianth, calyptra included, nat.
size.
30. Dehisecnt capsules; N. Taxilejeunea Colensoana showing valves not divided to the base, ×
10; O. Plagiochila × 7; P. Fossombronia × 7; Q. Metzgeria furcata with elaterophones × 10; R.
Asterella × 5.

Previous Figure | Table of Contents | Figure in Context | Next Figure

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Asterella elegans
(Elegant Asterella)

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Common Names
[ Back to top ]

Common Names in English:

Elegant Asterella

Description
[ Back to top ]

Physical Description

Habit: Nonvascular

Taxonomy
[ Back to top ]

Kingdom: Plantae ( ) - Plants

Phylum: Hepatophyta ( ) - Liverworts

Class: Hepatopsida ( )

Order: Marchantiales ( )

Family: Hydrophiloidea ( )

Genus: Asterella ( )

Specific epithet: elegans - (Spreng.) Trev.

Botanical name: - Asterella elegans (Spreng.) Trev.

Notes

Name Status: Accepted Name . Latest taxonomic scrutiny: 17-Oct-2001

Similar Species
[ Back to top ]

Members of the genus Asterella

There are approximately 102 species in this genus. Here are just 100 of them:

A. abyssinica · A. africana · A. aliena · A. alsophilae · A. angusta · A. aspidii · A. australis · A.

bachmannii · A. blumeana · A. bolanderi (Bolander's Asterella) · A. californica (California
Asterella) · A. capnoides · A. carnea · A. chichibuensis · A. clavuligera · A. confluens · A.
conocephala · A. conyzae · A. crustacea · A. dichaenoides · A. discoidea · A. dodonaei · A.
drummondii · A. echinella · A. elegans (Elegant Asterella) · A. erysiphoides · A. eupomatiae · A.
fissisquama · A. flexuosa · A. fumaginea · A. furcata · A. gardoquiae · A. gaultheria · A. gracilis
(Graceful Asterella) · A. graminicola · A. hellebori · A. himantia · A. infuscans · A. intricata · A.
karstenii · A. khasyana · A. lateralis · A. leptophylla · A. lindenbergiana (Lindenberg's Asterella)
· A. lindigii · A. liukiuensis · A. longiseta · A. ludwigii · A. macropoda · A. macularis · A.
manaosensis · A. marginata · A. megalospora · A. microtheca · A. missionum · A. muelleri · A.
multiplex · A. muscicola · A. mussuriensis · A. myricae · A. naumovii · A. nuda · A. oleina · A.
olivacea · A. opulenta · A. palmeri (Palmer's Asterella) · A. parmularia · A. patelloides · A.
paupercula · A. penicillata · A. peribebuyensis · A. phaeostroma · A. pseudocuticulosa · A.
pustulata · A. puttemansii · A. quscutae · A. rehmii · A. rubi · A. rubtzovii · A. rufoviolascens · A.
saccata (Asterella) · A. sanguinea · A. sanoana · A. schweinfurthii · A. setulosa · A.
stomatophora · A. subcuticulosa · A. subcyanea · A. subplana · A. tasmanica · A. tenella
(Asterella) · A. tenera · A. tenuissima · A. trichodea · A. valida · A. venosa · A. verruculosa · A.
virescens · A. wallichiana · A. whiteleggeana

Physical Description

Habit: Nonvascular

Taxonomy
[ Back to top ]

Domain: Eukaryota - Whittaker & Margulis,1978

Kingdom: Plantae - Haeckel, 1866 - Plants

Subkingdom: Viridaeplantae - Cavalier-Smith, 1981

Phylum: Bryophyta - A. Braun, in Ascherson, 1860 - Mosses

Subphylum: Hepaticae - (Auct.) Cavalier-Smith, 1998

Class: Hepaticopsida
Subclass: Marchantiidae

Order: Marchantiales

Suborder: Marchantiineae

Family: Aytoniaceae

Genus: Asterella

Specific epithet: palmeri - (Aust.) Underw.

Botanical name: - Asterella palmeri (Aust.) Underw.

Notes

Latest taxonomic scrutiny: 17-Oct-2001

Domain : Eukaryota Whittaker & Margulis,1978 - eukaryotes

Kerajaan : Plantae Haeckel, 1866

Subkerajaan : Viridaeplantae Cavalier-Smith, 1981 - Green Plants

Filum : Bryophyta A. Braun, in Ascherson, 1860 - Mosses

Subfilum : Hepaticae (Auct.) Cavalier-Smith, 1998

kelas : Marchantiopsida - Liverworts

Subkelas : Marchantiidae

Ordo : Marchantiales

Subordo : Marchantiineae

Suku : Aytoniaceae

Marga : Asterella

Domain : Eukaryota - Whittaker & Margulis,1978

Kerajaan : Plantae - Haeckel, 1866 - Plants

Subkerajaan : Viridaeplantae - Cavalier-Smith, 1981

Divisi : Bryophyta - A. Braun, in Ascherson, 1860 - Mosses

Subdivisi : Hepaticae (Auct.) Cavalier-Smith, 1998

Kelas : Hepaticopsida

Subkelas : Marchantiidae

Ordo : Marchantiales

Subordo : Marchantiineae

Suku : Aytoniaceae

Marga : Asterella

Jenis : Asterella gracilis (Web.) Underw.

Kerajaan : Plantae- Plants

Divisi : Hepatophyta - Liverworts

Kelas : Hepatopsida

Ordo : Marchantiales

Suku : Cercopoidea

Marga : Asterella

Jenis : Asterella tenella (L.) P.-Beauv.

Kerajaan : Plantae - Plants

Divisi : Hepatophyta - Liverworts

Kelas : Hepatopsida

Ordo : Marchantiales

Suku : Hydrophiloidea
Marga : Asterella

Jenis : - Asterella echinella

Kerajaan : Plantae - Plants

Divisi : Hepatophyta - Liverworts

Kelas : Hepatopsida

Ordo : Marchantiales

Suku : Hydrophiloidea

Marga : Asterella

Jenis : Asterella bolanderi (Aust.) Underw.

Kerajaan : Plantae - Plants

Divisi : Hepatophyta - Liverworts

Kelas : Hepatopsida

Ordo : Marchantiales

Suku : Hydrophiloidea

Marga : Asterella

Jenis : Asterella californica (Hampe) Underw.

Kerajaan : Plantae - Haeckel, 1866 - Plants

Subkerajaan : Viridaeplantae - Cavalier-Smith, 1981

Divisi : Bryophyta - A. Braun, in Ascherson, 1860 - Mosses

Subdivisi : Hepaticae - (Auct.) Cavalier-Smith, 1998

Kelas : Hepaticopsida

Subkelas : Marchantiidae

Ordo : Marchantiales

Subordo : Marchantiineae

Suku : Aytoniaceae

Marga : Asterella

Jenis : Asterella lindenbergiana (Corda) Lindb.

Kerajaan : Plantae - Plants

Divisi : Hepatophyta - Liverworts

Kelas : Hepatopsida

Ordo :Marchantiales

Suku : Cercopoidea

Marga : Asterella

Jenis : Asterella saccata (Wahlenb.) Evans

Rosa canina

Rosa canina juga dikenal dengan nama Brier hip atau brier rose atau dogberry,atau
dog rose,eglantine gall, hep tree, hip fruit, hip rose, hip tree, hop fruit, hogseed,
sweet brier, wild brier, Dog rose, rosal silvestre, roser de ca, rosa canina, , eglantier,
hecken-rose, hondsroos
dan witches' brier. Rosa canina tergolong dalam famili Rosaceae dan tipe dari
tanaman ini adalah semak belukar. Biasanya ditemukan pada daerah Eropa(Nova
Scotia, Virginia and Tennessee) dan Asia Barat. Bagian yang biasanya digunakan
sebagai pengobatan maupun bahan nutrisi adalah bagian buahnya yang berwarna
merah. Tanaman ini mampu menahana angi, namun tidak cocok hidup di daerah
maritim.
Sinonim : Rosa canina var. dumetorum Baker
Kingdom Plantae
Subkingdom Tracheobionta
Division Magnoliophyta
Class Magnoliopsida
Subclass Rosidae
Order Rosales
Family Rosaceae
Genus Rosa L.
Species Rosa canina L.

Rosa canina

Morfologi :
a. Daun (Folium)
Berdasarkan pertumbuhan daunnya, Rosa canina tergolong pada tumbuhan yang
berdaun tidak lengkap, yaitu hanya terdiri dari tangkai daun (petiolus) dan helaian
daun (lamina) saja, yang kemudian lazimnya disebut daun tidak bertangkai. Selain
itu, daun pada tanaman ini juga mempunyai alat-alat tambahan atau pelengkap yaitu
daun penumpu yang melekat pada kanan kiri pangkal tangkai daun (stipulae
adnatae) seperti pada jenis mawar pada umunya. Rosa canina termasuk dalam
golongan daun majemuk menyirip gasal. Berikut ini adalah keterangan mengenai
bagian daun dengan lebih rinci :
Tangkai daun(petiolus) : tangkai daunnya setengah lingkaran dan sisi atasnya
bergalur dangkal, berwarna hijau dan ditutupi bulu-bulu halus berwarna putih bahkan
ditumbuhi duri.
Helaian daun (lamina) : Sifat-sifat dari helaian daun sangat beragam, kondisi usia ari
tanaman juga mempengaruhi karakter helaian daun. Namun ada beberapa karakter
yang dapat dijadikan acuan, antara lain :
Bangun daun (circumscriptio): Berdasarkan letak bagian daun yang terlebar, daun
pada Rosa canina memiliki bagian yang terlebar pada bagian bawah tengah-tengah
helaian daun. Pangkal daunnya tidak bertoreh, dan bentuk bangunnya adalah
bangun bulat telur (ovatus).
Ujung daun (apex) : bentuk ujung daun pada Rosa canina runcing(acutus) yaitu pada
kedua tepi daun di kanan kiri ibu tulang sedikit demi sedikit menuju ke atas dan
pertemuannya pada puncak daun membentuk suatu sudut lancip(lebih kecil dari
900).
Pangkal daun (basis) : Pangkal daunnya tidak pernah bertemu namun terpisah oleh
pangkal ibu tulang/ ujung tangkai daun. Pangkal daunnya membulat (rotudantus),
yaitu pada daun-daun bangun bulat, jorong, dan bulat telur.
Susunan tulang daun (nervatio atau venatio) : Tulang-tulang daun menurut besar
kecilnya dibedakan dalam 3 macam, yaitu
Ibu tulang(costa) : bentuknya simetrik.
Tulang-tulang cabang : tulang cabangnya ada yang tingkat 1, yaitu tulang cabang
yang langsung berasal dari ibu tulang. Ada pula tulang cabang tingkat 2 yaitu tulang
cabangnya berasal dari tulang cabang tingkat 1.
Urat-urat daun : tulang cabang dapat mencapai daun.
Melihat arah tulang-tulang cabang yang besar pada helaian daun-daun, Rosa canina
pertulangannya adalah bertulang menyirip (pennivernis), yaitu daun ini mempunyai
satu ibu tulang yang berjalan dari pangkal ke ujung, dan merupakan terusan tangkai
daun. Dari ibu tulang ini ke samping keluar tulang-tulang cabang sehingga
susunannya mengingatkan kita kepada susunan sirip-sirip pada ikan, oleh sebab itu
dinamakan pertulangan menyirip. Dari sinilah kita dapat tahu bahwa Rosa canina
adalah tanaman dikotil sebab salah satu ciridari tanaman dikotil adalah pertulangan
daunnya yang menyirip.
Tepi daun (margo) : bergerigi(serratus), yaitu sinus dan angulus sama lancip. Tipe
margo ini tergolong dalam tepi daun dengan toreh merdeka di mana bentuk
torehnya tidak mempengaruhi bentuk.
Daging daun (intervenium) : seperti kertas (papyraceus atau chartaceus) tipis namun
cukup tegar.
Keadaan permukaan daun : berbulu(pilosus),
Warna daun : pada bagian atas berwarna hijau tua, pada bagian bawah juga
berwarna hijau namun agak lebih muda.
Susunan daun pada tanaman Rosa canina ini maka dapat digolongkan pada tanaman
daun majemuk menyirip gasal (impapiripinnatus), sebab jika ditinjau dari jumlah
anak daunnya akan kita dapati bilangan yang benar-benar gasal pada anak daun
yang berpasangan, sedagkan pada di ujung ibu tangkai terdapat anak daun yang
tersendiri (dengan ukuran yang lebih besar daripada yang lainnya). Lihat gambar di
bawah ini:

Gambar daun Rosa canina

b. BATANG (Caulis)
Batangnya termasuk batang yang berkayu (lignosus) yaitu batangnya keras dan
kuat, namun bentuk dari tanaman ini adalh semak(frutices). Berikut ini adalah
keterangan lebih rinci mengenai batang:
Bentuk batang : bentuk batangnya bulat (teres).
Permukaan batang : beralur(sulcatus), pada batang terdapat alur-alur yang jelas.
Arah tumbuh batang : memanjat (scandens) sebab batang dapat tumbuh ke atas
dengan menggunakan penunjang. Penunjang pada tanaman Rosa canina ini, adalah
duri. Lihat gambar di bawah ini

Pertumbuhan memanjat dari Rosa canina

Percabangan pada batang : tergolong ke dalam percabangan simpodial, di mana
batang pokok sukar ditemukan karena dalam perkembangan selanjutnya,
menghentikan pertumbuhannya atau kalah besar dan kalah cepat pertumbuhannya.
Arah tumbuh cabang : tegak (fastigiatus) sebab sudut antara batang dan cabang
amat kecil, sehingga arah tumbuh cabang hanya pada pangkalnya saja sedikit
serong ke atas, tetapi selanjutnya hamper sejajar dengan batang pokoknya.
Warna batang : coklat tua pada yang sudah tua.

Gambar batang Rosa canina yang berduri

c. BUAH (fructus)
Buahnya adalah buah sejati, yaitu buah sejati ganda di mana terjadi dari satu bunga
dengan beberapa bakal buah yang bebas satu sama lain, dan masing-masing bakal
buah menjadi satu buah namun tetap berkumpul di sekitar tempat bunga itu tumbuh.
Berdasarkan sifat ini, maka buahnya tergolong dalam buah sejati kurung ganda,
yaitu dalam badan yang berasal dari dasar bunganya yang berbentuk periuk terdapat
banyak buah-buah kurung. Buahnya tumbuh pada musim gugur.

Gambar buah Rosa canina yang kaya akan sumber vitamin C, inset bunga sebelah
atas kiri menunjukkan awal mula adanya buah tersebut.
d. BIJI (semen)
Pada bagian biji ini terdapat bagian yang khas pada kulit biji(spermodemis), yaitu
adanya bulu (coma), maka pada pengkonsumsian buah ini harus berhati-hati karena
adanya bulu-bulu di sekitar bijinya, sebab dapat menyebabkan iritasi pada bagian
pencernaan. Pada bagian lembaga(embryo). Dilihat dari bijinya maka tanaman Rosa
canina ini tergolong ke dalam tanaman dikotil.

e. BUNGA (flos)
Pada tanaman ini, bunganya tergolong dalam bunga majemuk tak
berbatas(inflorescentia racemosa atau inflorescentia botryoides atau inflorescentia
centripetala), sebab ibu tangkainya dapat tumbuh terus, dengan cabang-cabang
yang dapat bercabang lagi atau tidak, dan susunannya “acropetal”(semakin muda
semakin dekat dengan ujung ibu tangkai) Dalam golongan ini, pertumbuhan
bunganya tergolong lagi ke dalam bentuk tandan (racemus atau botrys), sebab ibu
tangkainya bercabang dan cabangnya masing-masing mendukung satu bunga pada
ujungnya. Bunga Rosa canina ini tergolong bunga berkelamin dua(hermaphroditus)
dengan alat kelamin 2 yaitu putik(pistillum) dan benang sari (stamen). Bunganya
tergolong ke dalam bunga lengkap atau bunga sempurna karena terdiri atas
lingkaran daun-daun kelopak, lingkaran daun mahkota, lingkaran benang-benang sari
dan lingkaran daun-daun buah. Berdasarkan bidang simetri pada mahkota bunganya,
maka dapat pula dilihat pada gambar, bunga ini memiliki banyak bidang simetri
saja(polysimetris atau actinomorphus). Tajuk bunganya beraturan (regularis) dan
bentuknya adalah bintang(rotatus atau stellatus). Susunan benang sarinya tampak
seperti duduk di atas kelopak(calyciflorae).Berdasarkan jumlahnya, maka benang
sarinya masuk ke dalam golongan benang sari banyak. Putiknya merupakan putik
tunggal (simplex) putik hanya tersusun atas sehelai daun buah saja. Menurut
letaknya terhadap dasar bunga, maka bunga ini tergolong ke dalam bakal buah bakal
buah setengah tenggelam (hemi inferus), seba letak bakal buahnya duduk pada
dasar bunga yang cekung, sehingga tempat duduknya bakal buah lebih rendah
daripada tepi dasar bunga, dan sebagian dinding bakal buah itu berlekatan dengan
dasar bunga yang berbentuk mangkuk piala.

Gambar bunga Rosa canina

F. AKAR (radix)
Akar pada tanaman ini tergolong dalam sistem akar tunggang sebab akar lembaga
tumbuh terus menjadi akar pokok yang bercabang-cabang menjadi akar yang lebih
kecil. Akar tunggang yang terdapat pada tanaman ini adalah bentuk akar tunggang
dengan keadaan berbentuk sebagai tombak(fusiformis).

ANatomi
Pada bagian tanaman yaitu pada batang yang dipotong membujur, dinamai sebagai
poros cincin dan semi poros cincin. Pori-porinya dibatasi oleh jarak yang lumayan
besar. Parenkimanya apotracheal dan tersebar. Pada jaringan dasarnya cukup tebal.
Terdapat cambium dalam jumlah kecil namun pada deret yang cukup tebal. Lihat
gambar ini.

Jika dipotong melintang, berbentuk piringan, terlihat bagian yang berbentuk spiral.
Mengandung kristal berbentuk prismatic di dalamnya. Terdapat trakeid namun tidak
ada libiform. Lihat gambar di bawah:

Jika dipotong secara tangensial, tampak ada 2 tipe yaitu sel berbentuk oval dan yang
majemuk. Lihat gambar di bawah :

Kandungan kimia dan fisiologi

Pada buah yaitu :

ASAM ASKORBAT 7,400 - 25,000 ppm

ALPHA-TOCOPHEROL 41 - 204 ppm;
ALUMINUM 5 - 12 ppm
BETA-CAROTENE 25 - 62 ppm
BETULIN
BORON 8 - 22 ppm
BROMIN 2 - 5 ppm
CADMIUM 0.03 - 0.075 ppm
CALCIUM 3,500 - 8,750 ppm
ASAM KAPRIC
KARBOHIDRAT 387,600 - 779,000 ppm
CATECHIN
KLORIN313 ppm
KROMIUM0.03 - 0.07 ppm
ASAM SITRAT 15,000 ppm
COBALT 0.01 - 0.025 ppm
COPPER 1.8 - 36 ppm
D-ABSCISSIN
EO Seed 2,000 - 3,000 ppm
EPICATECHIN
LEMAK 17,000 - 26,000 ppm
SERAT 300,000 ppm
FLAVONOIDS 100 - 3,500 ppm
FLUORINE0.6 - 1.5 ppm
FRUKTOSA

GALLOCATECHIN
GLUKOSA
INVERT-SUGARS 100,000 - 137,000 ppm
IRON 5 - 13 ppm
ISOQUERCITRIN
KAEMPFEROL-3(P-COUMAROYLGLUCOSIDE)
KAEMPFEROL-3-GLUCOSIDE
LEAD 0.16 - 0.4 ppm
LEUCOANTHOCYANINS
LEUCOPAENIDIN
LEUCOROSINIDIN
LYCOPENE
MAGNESIUM 560 - 2,090 ppm
MALIC-ACID 5,000 - 62,200 ppm
MANGANESE 12 - 40 ppm
MERCURY
MOLYBDENUM
NIACIN
NICKEL 0.4 - 1 ppm
NITROGEN 3,600 - 9,000 ppm
PECTIN 110,000 ppm
PHOSPHORUS 560 - 5,180 ppm
POTASSIUM 5,843 - 21,000 ppm
PROTEIN 72,300 - 133,000 ppm
RIBOFLAVIN 7.2 ppm
RUBIDIUM 10 - 25 ppm
RUBIXANTHIN
SELENIUM 0.002 - 0.005 ppm
SILICON 10 - 25 ppm
SODIUM 2,930 - 4,600 ppm
SUCCINIC-ACID SUCROSE 6,000 - 32,100 ppm
SUGARS 116,000 - 156,000 ppm
SULFUR 400 - 1,000 ppm
TANNIN 20,000 - 27,000 ppm
TARAXAXANTHIN
THIAMIN 3.8 ppm
TIN 22 ppM
VANILLIN
WATER 502,000 - 913,000 ppm
XANTHOPHYLL
ZEAXANTHIN

Pada akar yaitu :

TANNINS 15,000 ppm
TRITERPENES 5,200 ppm
PYROGALLOTANNINS 15,000 ppm

Pada biji yaitu :

EO 2,000 - 3,000 ppm
LECITHIN
ASAM LINOLEIC 43,200 - 58,320 ppm
ASAM ASETAT
ASAM LINOLENIC 25,600 - 34,560 ppm
TOCOPHEROLS 660 ppm

Pada daun yaitu :

HISTONE
TANNINS 100,000 - 260,000 ppm

Tanaman ini melakukan sintesis C3 untuk membentuk glukosa. Diawali dengan

fiksasi CO2, yaitu menggabungkan CO2 dengan sebuah molekul akseptor karbon.
Akan tetapi di dalam sintesis C3, CO2 difiksasi ke gula berkarbon 5 yaitu ribulosa
bifosfat (RuBP) oleh enzim karboksilase RuBP (rubisko). Molekul berbentuk 6 yang
tidak stabil dan segera terpisah menjadi 2 molekul yaitu fosfogliserat(PGA). Molekul
PGA merupakan karbohidrat stabil berkarbon 3, yang pertama kali terbentuk,
sehingga cara tersebut dinamakan sintesis C3 .Molekul PGA bukan molekul berenergi
tinggi. Dua molekul PGA mengandungenergi yang lebih kecil dibandingkan denagn 1
molekul RuBP. Hal tersebut menjelaskan alas an fiksasi CO2 berlangsung secara
spontan dan tidak memerlukan energi dari reaksi cahaya. Untuk mensintesis molekul
berenergi tinggi energi dan electron dari ATP maupun NADPH hasil reaksi terang dan
digunakan untuk mereduksi tiap PGA menjadi fosfogliseraldehida (PGAL). Dua
molekul PGAL dapat membentuk 1 molekul glukosa. Siklus Calvin telah lengkap bila
pembentukan glukosa disertai dengan regenerasi RuBP. Satu molekul CO2 yang
tercampur menjadi 6 molekul CO2. Ketika enam molekul CO2 bergabung dengan
enam molekul RuBP dihasilkan satu glukosa dan 6 RuBP sehingga siklus dapat
dimulai lagi.
Kegunaan

Rosa canina biasanya digunakan untuk obat diuretic yang aman bagi ginjal. Selain
itu, buahnya baik untuk mengobati penyakit pada ginjal dan radang kantung kemih,
flu,diare selain itu juga dapat digunakan untuk demam, astringen, tonik dan pencita
rasa pada teh. Sebagai tambahan, pada masa perang Dunia ke 2, buah Rosa canina
menggantikan sumber vitamin C pada masa itu.