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Moss (Bryophyte) (蘚苔類)

Mount Usu / Sarobetsu post-mined peatland
From left: Crater basin in 1986 and 2006. Cottongrass / Daylily

A rolling stone gathers no moss

Embryophyta (Cronquist)

= mosses (bryophyta) + ferns (pteridophyta) + seed plants (spermatophyta)
Moss (s.l.): = bryophytes
Moss (s.s.): bryophytes = mosses + hornworts + liverworts*

= non-vascular terrestrial plants: no vascular tissues
→ small due to not having vascular tissues
5000-25000 species

Plagiochima: 7 species in Hokkaido and 8 species in Kyushu
Apomixis (weak speciation) → population maintenance
  1. chloroplast segments: chlorophyll a, b, carotine α, β, xanthophyll, lutein
  2. starch
  3. cellulose
  4. sperm = whiplash-typed flagellatum whiplash
  5. thallophyte ↔ cormophyte
moss cycle
Sproe(n) → Protonema(n) → Juvenile(n)
  ↑R.D.                       ↓
Sporophyte(2n) → Mature plant (= Gametophyta, n):
                  ↓  ♂ ♀ reproductive organ <─────┐
      sexual reproduction (G)  ↓ vegetative reproduction │
                           Mature plant ─────────┘

Fig. Life cycle of moss (→ fern)

moss cycle
Sporophytes(2n) grow on gametophytes(n)
→ nutrients are abosorbed by feet (足) on sporophytes
→ 2n parasitizes n = moss-typed life history
[ species | Alaska ( feather moss )| plant taxonomy ( 植物分類学 )]


Musci: n = 5-60(120?), commonly having 10, 11 or 12
Hepaticae: n = 4-32, commonly having 9

n of 40 species in Barrow (Alaska)

37 spp = 9
2 spp = 18 diploid (9 × 2)
1 spp = 8 haploid (9 - 1)

n of 50 species in the Magellan Strait

n = 9 for all the speces
Morphologically diverse
Asexual reproduciton for most species

Anthocerotae: n = 5

n = 5 for 19 species measured The number of chromosome is extremely small of the terrestrial plants

Takakia (ナンジャモンジャゴケ), liverworts): n = 4 (2U + 2J Takakia

Two types of chromosomes, U and J
Only Takakia has n = 4 (Re-examining the reports from the other species)

on the evolution of chromosome numbers
n = 4 → 8 → 8 + 1 → 18 → ···

the proposal was interpreted as negative at first

Takakia having n = 5 was discovered
The two types are morphologically different, e.g., cell wall that is thicker in the type n = 5
→ Two species were established

T. lepidoziodes: n = 4
T. ceratophylla: n = 5

Discovered that Takakia having n = 4 showed morphologically n = 5

T. lepidozoides: n = 5 = 2V + 2J + F
T. ceratophylla: n = 5 = 2V + 3J

Takakia F chromosome was considered to be derived from the fragment of J chromosome
Takakia: the direction of evolution is likely to be decrease in chromosome numbers
The stage having F chromosome shows the morphological traits of T. lepidozioides, although the n is 5

Musci or bryophyta (蘚類)

Class Musci (蘚類綱): Three subclasses =
= Sphagnidae + Andreaeidae (クロゴケ) + Brydae (マゴケ)
leafy moss gametophytes develop from a protonema
a moss sporophyte consists of a capsule, seta and foot
Protonema: a filament of n cells that grows from a spore and develops into leafy moss gametophytes in mosses
Capsule: portion of the bryophyte sporophyte in which spores are produced

[peat (泥炭)]

Subclass Sphagnidae (ミズゴケ), peat moss


Sphagnum L. (ミズゴケ)
peat moss
Fig. Morphological traits.
Hyaline cell (透明細胞): non-photosynthetic, porous cells found in Sphagnum leaves
S. acutifolium Ehrh. ex Schrad. var. flavescens Warnst. (S. acutifolium Scop.* former name)
S. acutum Warnst. var. hakusanense Warnst.ハクサンミズゴケ
S. angustifolium (C.Jens.) C.Jens.アオモリミズゴケ*
S. calymmatophyllum Warnst. et Card. コバノミズゴケ
S. connectans Warnst. et Card.イトナシミズゴケ
S. cuspidatum var. submersum Schimp.シワミズゴケ
S. dicladum Warnst.ヤマミズゴケ
S. drepanocladum Warnst. var. latilimbatum Warnst.ヘリトリミズゴケ
S. fallax Klinngr.
S. flexuosum Dozy et Molk.
S. imbricatum Hornsch. ex Russ.フナガタミズゴケ (クシバミズゴケ*)

var. affine (Ren. et Card.) Warnst.ハリマミズゴケ)

S. incertum Warnst. et Card.ミネミズゴケ
S. jensenii Lindb.シナノミズゴケ*
S. kushiroense H.Suzukiクシロミズゴケ
S. lindbergii Schimp. ex Lindb.フサバミズゴケ
S. majus (Russ.) C. Jens.
S. magellanicum Bird. (ムラサキミズゴケ):
Sphagnum1 Sphagnum2
[1/2] in Poker Flat Research Range, interior Alaska, on August 12 2012.
S. microporum Warnst. ex Card. コアナミズゴケ: aquatic, curved branches, small pores at the backside of transparent cell
S. obtusum Warnst.
S. pallens Warnst. et Card. シロミズゴケ
S. palustre (オオミズゴケ)

var. pallescens (Warnst.) Sak.ウスアオミズゴケ
var. virescens (Russ.) Sak.アオオオミズゴケ
Fig. S. palustre

S. papillosum Lindb. イボミズゴケ
Distribution: boreal regions in the Northern Hemisphere (common in a post-mined peatland, Sarobetsu Mire, northern Hokkaido)
Sphagnum1 Sphagnum2
[1/2] along the shore of Echigo Pond in Ebetsu City on October 8 2017.

var. confertum Lindb.フトミズゴケ
var. riparium Grav.

S. platyphyllum (Lindb.) Warnst.
S. plumulosum Roellワラミズゴケ*
S. pulchrum (Lindb. ex Braithw.) Warnst. ウツクシミズゴケ
S. quinquefarium (Lindb. ex Braithw.) Warnst.ゴレツミズゴケ
S. recurvum P. Beauv.サンカクミズゴケ
S. riparium Angstr.サケバミズゴケ*

var. coryphaceum Russ.セイタカミズゴケ)

S. rubellum Wils.ウスベニミズゴケ(アカミズゴケ*)
S. russowii Warnst.
S. septatum Warnst.フシミズゴケ
S. subacutifolium Schimp.スギバミズゴケモドキ
S. subobesum Warnst.シタミズゴケ
S. teres (Schimp.) Angstr. ex C.Hartm.ホソミズゴケ *
S. tosaense Warnst. トサミズゴケ
S. warnstorfii Russ.ワルンストロフミズゴケ
Sphagnum sheep

Sphagnum sheep

Residual Sphagnum hummocks that survived fire activity
Sphagnum sheep provides safe sites for the seedling establishment of Picea mariana, and contributes micro-climate changes, such as albedo.

Subclass Andreaeidae (クロゴケ亜綱)

Subclass Brydae (マゴケ亜綱)

Subclass Buxbaumidae (キセルゴケ亜綱)

Subclass Polytrichidae (スギゴケ亜綱)

Order Polytrichales (スギゴケ)

Polytrichacae (スギゴケ)

Order Dawsoniales

Polytrichum1 Polytrichum2
Polytrichum juniperinum Willd. ex Hedw. (juniper haircap or juniper polytrichum moss, スギゴケ)
[1/2] in a post-mined peatland, Sarobetsu mire on May 22 2016.
Polytrichum1 Polytrichum2
Polytrichum commune Hedw. (Common haircap moss, ウマスギゴケ)
[1/2] at Poker Flat that was burned in the summer of 2004, Fairbanks, AK, on August 10 2012. This species is widespread on the site , as a pioneer on succession (Tsuyuzaki et al. 2009)
Polytrichum1 Polytrichum2
Polytrichum piliferum Hedw. (Bristly haircap, ハリスギゴケ)
[1/2] clsoe to an area damaged by the volcnaic eruptions in 1996 on the summit of Mount Koma, southern Hokkaido, on July 7 2013. n = 7. The annual fluctuation of plant cover was highly variable in plots established on the mountain.

Hepaticae (苔類), liverworts


Fig. Thallose and leafy shoot

L: lateral merophyte, V: ventral merophyte

(Inoue 1975)

Class Hepaticae (苔類綱)
Subclass Calobryidae (コマチゴケ亜綱)
Order Takakiales
Order Calobryales Haplomitriaceae Ded. (コマチゴケ)
Subclass Jungermanniidae (ウロコゴケ亜綱)
Order Metzgeriales (Anacrogynae) フタマタゴケ
lack air chambers, pores, ventral scales
gametangia and sporophytes are sessile on the gametophyte
elevation of the capsule (sporangium) is accomplished by elongation of the seta (stalk) of the sporophyte
sporangia contain elaters
Order Jungermanniales (Acrogynae) (ウロコゴケ, leafy liverworts)
dorsiventral growth habit
two rows of lateral leaves and one row of ventral leaves (amphigastria)
Subclass Marchantiidae (ゼニゴケ亜綱)
Marchantiaceae (ゼニゴケ)
Marchantia L. (ゼニゴケ), merged with Bucegia Radian and Preissia Corda
M. polymorpha L. (ゼニゴケ, common liverwort)
female plant with archegoniophores / Male plant with antheridiophores
gemmae cups
dorsal surface of thallus showing air chamber outlines and pores
Marchantia Marchantia Marchantia
Fig. (left/center) M. polymorpha: at a roadside in the city of Sapporo (N22/W2, East Ward) on October 26 2009. This species is common in various habitats, e.g., volcanoes and post-fire forest, as a pioneer on succession. (right) antheridiophore at a roadside in N27/E2, Sapporo, on May 1 2018.
Ricciaceae (ウキゴケ):
Riccia (ウキゴケ): most species are terrestrial →

R. fluitans: amphibious, by large air chambers in the thallus

Ricciocarpus (イチョウウキゴケ)

antheridia and archegonia - (monoecious)
antheridia - protandrous

Anthocerotae (ツノゴケ), hornworts

the sporophyte is shaped like a tapered horn
the sporophyte has an intercalary meristem, so can grow indeterminately the thallus has stoma-like structures, the only known occurrence in a gametophyte
each photosynthetic cell contains a single chloroplast
archegonia are embedded in the thallus and in contact with surrounding vegetative cells
cavities are filled with mucilage containing Nostoc

Species (種)

Plagiomnium1 Plagiomnium2
Bryhnia novae-angliae (Sull. et Lesq.) Grout (ヤノネゴケ)
Distribution: temperate and cool-temperate regions in the Northern Hemisphere
Habitat: moist and wet habitats, occasionally in water
[1/2] in the Furano Experimental Forest of the University of Tokyo, central Hokkaido, on June 28 2017
Plagiomnium1 Plagiomnium2
Plagiomnium acutum (Lindb.) T. J. Kop. (コツボゴケ)
Distribution: widespread in eastern Eurasia, including the whole of Japan
Habitat: on soil and rock
[1/2] near a transplanted forest of Fraxinus mandshurica in the Experimental Forest of the University of Tokyo, Furano, central Hokkaido, on June 26 2017.
Glyphomitrium1 Glyphomitrium2
Glyphomitrium humillimum (Mitt.) Card. (サヤゴケ)
Distribution: widespread in the East Asia, including the whole of Japan
Habitat: on bark
[1/2] in front of Sankai-no-taki (三階の滝, Triple-floor Falls), central Hokkaido, on June 26 2017.
Climacium1 Pohlia nutans2
Pohlia nutans (Hedw.) Lindb. (ヘチマゴケ)
[1/2] on a concrete fence near a post office, N14/W5, North Ward, Sapporo, on May 8 2013.
Climacium1 Climacium2
Climacium japonicum Lindberg (コウヤノマンネングサ)
[1/2] at Tomakomai Experimental Forest on August 23 2012. A few picture books describe that ths plant is beautiful.
Bryum1 Bryum2
Bryum argenteum Hedw. (ギンゴケ)
Distribution: cosmopolitan, including antarctic
Chromosome number: 2n = 10
[1/2] on an unpaved parking lot, N27/E2, East Ward, Sapporo, on November 17 2015.
Bryum1 Bryum2
Ceratodon purpureus (Hedw.) Brid. (ヤノウエノアカゴケ)
Distribution: cosmopolitan (Ceratodon consists of a few species but is one species in Japan)
Habitat: disturbed sites (becoming common soon after wildfire and eruption)
[1/2] in front of the Center for Advanced Medical Imaging, Sendai Welfare Hospital, northern Japan, on March 23 2016.
Bryum1 Bryum2
Bartramia pomiformis Hedw. (タマゴケ)
Distribution: broadly distributed in the Northern Hemisphere (east Asia - Siberia - Europe - North America)
Habitat: developing colonies on soils and rocks
Chromosome number: n = 8
[1/2] on Oshima Island in Matsushima Islands, northern Honshu, on March 21 2016.
Rhacomitirum1 Rhacomitirum2
Racomitrium lanuginosum (Hedw.) Brid. (シモフリゴケ)
Distribution: widespread in the world
Habitat: developing large carpets on soils and rocks
[1/2] on the southwestern slope of Mount Koma, southern Hokkaido, on July 9 2017.


Moss flora list on Poker Flat (after the 2004 wildfire)
The specimens are stored in SAPS. Thank Dr. Higuhi for the identification!
species checklist in Alaska
  • Species (Japanese name, English name): Remarks
  • Aulacomnium palustre (Hedw.) Schwagr (オオヒモゴケ)
  • Aulacomnium turgidum (Wahl.) Schwagr. (フトヒモゴケ)
  • Ceratodon purpureus (Hedw.) Brid. (ヤノウエノアカゴケ, fire moss): on burned ground suface and dead Sphagnum moss
  • Hylocomium splendens (Hedw.) B. S. G. (イワダレゴケ, splendid feather moss)
  • Marchantia polymorpha L. (ゼニゴケ, common liverwort)
  • Pleurozium schreberi (Brid.) Mitt. (タチハイゴケ, red-stemmed feather moss): forest floor in subalpine zoenes (Japan)
  • Polytrichum commune Hedw. (ウマスギゴケ, common haircap moss)
  • Polytrichum juniperinum Willd. ex Hedw. (スギゴケ, juniper haircap moss)
  • Sphagnum lenense H. Lindb. (コフサバミズゴケ)
  • Sphagnum capillifolium (Ehrh.) Hedw. (スギハミズゴケ)
  • Sphagnum cuspidatum Ehrh. ex. Hoffm. (ハリミズゴケ, toothed peat moss)
  • Sphagnum fuscum (Schimp.) Klinggr. (チャミズゴケ, rusty peat moss)
  • Sphagnum lindbergii Schimp. (フサバミズゴケ)
  • Sphagnum magellanicum Brid. (ムラサキミズゴケ, Magellan's peat moss)

Feather moss

  1. feather moss, boreal forest moss, or plume moss
    Ptilium crista-castrensis (Syn. Hypnum crista-castrensis) [dachougoke in Japanese, ダチョウゴケ]
    sparse thallus
    → The Linnean Collections: Hypnum crista-castrensis
  2. splendid feather moss, fern moss, or step moss
    Hylocomium splendens (Hedw.) Schimp. [iwadaregoke, イワダレゴケ]
    dense thallus
  3. feather moss (UK), or brachythecium moss
    Brachythecium albicans (Hedw.) Schimp.
  4. common feather moss (UK), slender beaked moss (Canada)
    Eurhynchium praelongum (Hedw.) B.S.G.
P. crista-castrensis and H. splendens are common in Picea mariana forests.
Cf. Red-stemmed feather moss (or, Boreal feather moss)

= Pleurozium schreberi [tachihaigoke, タチハイゴケ]