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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 species
Morphologically diverse
Asexual reproduction 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
T. lepidozioides Hattori et Inoue ナンジャモンジャゴケ

n = 4 (2V + 2J)


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

Hypothesis on the evolution of chromosome numbers

n = 4 → 8 → 8 + 1 → 18 → …
the proposal was interpreted as negative at first

T. ceratophylla: discovered n = 5
The two morphological types, e.g., cell wall thicker in the type n = 5

Two species were established
T. lepidoziodes, n = 4 and 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 or 2V + 2J + v

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
Capitulum (頭部, pl. -a): compact clusters of branches on the top
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), developing divergent branch (開出枝) and pendulous branch (下垂枝)

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. squarrosum Crome (ウロコミズゴケ, spiky bog-moss)
Sphagnum1 Sphagnum2
[1] in a forest (Symbiosis Forest) near Nopporo Forest Park on May 5 2020. [2] on a wetland forest in the Shibecha Experimental Forest of Kyoto University, eastern Hokkaido, on June 27 2019.
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)

Andreaeaceae Dumort. (クロゴケ)
Acroschisma (Hook. f. et Wilson) Lindl.

A. andensis Spruce, southern America

Andreaea Hedw. (クロゴケ)
Small, delicate acrocarpous mosses
Forming dark brown or reddish cushions on wet siliceous rocks

Subclass Brydae (マゴケ亜綱)

Order Tetraphidales (ヨツバゴケ)

Order Fissidentales (ホウオウゴケ)

Order Archidiales (ツチゴケ)

Order Dicranales (シッポゴケ)

Ditrichanceae (キンシゴケ)
Ceratodon purpureus (Hedw.) Brid. (ヤノウエノアカゴケ)
Bryum1 Bryum2
Distribution: cosmopolitan (Ceratodon consists of a few species but is one species in Japan)
Habitat: disturbed sites (becoming common soon after wildfire and eruption)

Order Pottiales (センボンゴケ)

Pottiaceae (センボンゴケ)
25 species in Japan
Barbula Hedw. (ネジクチゴケ)
B. unguiculata Hedw. (ネジクチゴケ), n = 13
Barbula unguiculata Barbula unguiculata
in a house garden, N14/W3, North Ward, Sapporo, on April 13 2018.

Order Grimmiales (ギボウシゴケ)

Grimmiaceae (ギボウシゴケ)
Aligrimmia R. S. Williams
Bucklandiella Roiv.
Codriophorus P. Beauv.
Coscinodon Spreng.
Coscinodontella R. S. Williams
Dryptodon Brid.
Gasterogrimmia (Schimp.) Buyss.
Grimmia Hedw.
Glyphomitrium Bridel
G. humillimum (Mitt.) Card. (サヤゴケ)
Glyphomitrium Glyphomitrium Distribution: widespread in the East Asia, including the whole of Japan
Habitat: on bark
Guembelia Hampe
Indusiella Broth. et Müll. Hal.
Jaffueliobryum Thér.
Leucoperichaetium Magill
Niphotrichum (Bedn.-Ochyra) Bedn.-Ochyra et Ochyra
Orthogrimmia (Schimp.) Ochyra et Żarnowiec
Racomitrium Brid. (シモフリゴケ)
R. lanuginosum (Hedw.) Brid. (シモフリゴケ)
Racomitrium Racomitrium
Distribution: widespread in the world (incluidng Mount Koma)
Habitat: developing large carpets on soils and rocks
Schistidium Bruch et Schimp.
Scouleria Hook.
Streptocolea (I. Hagen) Ochyra et Żarnowiec
Tridontium Hook. f.

Order Funariales (ヒョウタンゴケ)

Order Schistotegales (ヒカリゴケ)

Order Eubryales (ホンマゴケ)

Bryaceae (カサゴケ)
Bryum Hedw. (ハリガネゴケ)
Bryum argenteum Hedw. (ギンゴケ)
Distribution: cosmopolitan, including antarctic
Chromosome number: 2n = 10
Bryum Bryum
Polia Hedw. (ヘチマゴケ)
P.E nutans (Hedw.) Lindb. (ヘチマゴケ)
Pohlia nutans Pohlia nutans
on a concrete fence near a post office, N14/W5, North Ward, Sapporo, on May 8 2013.
P. flexuosa Hook. (ケヘチマゴケ)
Pohlia nutans Pohlia nutans
Mniaceae Schwägr. (チョウチンゴケ)
Plagiomnium T. J. Koponen (ツルチョウチンゴケ)
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
Plagiomnium Plagiomnium
Pohlia Hedw. (ヘチマゴケ)
Bartramiaceae Schwägrichen (タマゴケ)
Bartramia Hedw. (タマゴケ)
B. 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
Bryum Bryum
on Oshima Island in Matsushima Islands, northern Honshu, on March 21 2016.

Order Orthotrichales (タチヒダゴケ)

controversial group

Order Isobryales (イヌマゴケ)

Climaciaceae (コウヤノマンネングサ)
C. japonicum Lindberg (コウヤノマンネングサ)
Chromosome number: n = 11
Climacium Climacium
in Tomakomai Experimental Forest on August 23 2012. A few picture books describe that ths plant is beautiful.

Order Hookeriales (アブラゴケ)

Order Hypnobryales (シトネゴケ)

Theliaceae (ヒゲゴケ)
Fabroniaceae (コゴメゴケ)
Leskeaceae (ウスグロゴケ)
Thuidiaceae (シノブゴケ)
Abietinella C. Muell. (モミシノブゴケ)
Anomodon Hook. et Tayl. (キヌイトゴケ)
Boulaya Card. (チャボスズゴケ)
Clapodium (Lesq. et James) Ren. et Card. (ハリゴケ)
Haplocladium (コバノキヌゴケ)
H. angustifolium (Hampe et Mull. Hal.) Broth. (ノミハニワゴケ)
Haplocladium Haplocladium
in front of GSEES, HU, on April 12 2019.
Miyabea Broth. (ミヤブゴケ)
Thuidium B.S.G. (シノブゴケ)
Amblystegiaceae (ヤナギゴケ)
Brachytheciaceae (アオギヌゴケ)
Brachytecium (ネズミノオゴケ)
Bryhnia Kaurin
B. novae-angliae (Sull. et Lesq.) Grout (ヤノネゴケ)
Plagiomnium Plagiomnium
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 Homalothemium
Entodontaceae (ツヤゴケ)
Plagiotheciaceae (サナダゴケ)
Sematophyllaceae (ハシボソゴケ)
Hypnaceae (ハイゴケ)
Rhytidiaceae (フサゴケ)
Hylocomiaceae (イワダレゴケ)

Subclass Buxbaumiidae (キセルゴケ亜綱)

Order Buxbaumiales (キセルゴケ)

Buxbaumiaceae (キセルゴケ)
pioneer species on succession
Buxbaumia Hedw. (キセルゴケ), bug moss
Diphyscium D. Mohr (イクビゴケ)
Theriotia (クマノゴケ)

Subclass Polytrichidae (スギゴケ亜綱)

Order Polytrichales (スギゴケ)

Polytrichacae (スギゴケ)
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)

Other common names: common haircap, great golden maidenhair, great goldilocks, common haircap moss or common hair moss

Distribution: throughout temperate and boreal regions in the Northern Hemisphere and several Pacific Islands including New Zealand, and in Australia

Habitat: in bogs, wet heathland and along forest streams

xeromorphic adaptation

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.

Order Dawsoniales


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)
Plagiochilaceae Müll. Frib. (ハネゴケ)
characterized by perianths laterally compressed, bilabiate and wide-mouthed
sex organs not in specialized positions
capsule walls many-stratose; and seta thick
1600 species in the world, many endemic species

all species in Hawaii are endemic

Acrochila: A. caledonica (Steph.) Inoue
Chiastocaulon dendroides (Nees) Carl
Plagiochila: P. spinulosa (Dicks.) Dum., P. wakawana Steph., P. subopposita Steph.
Plagiochilion: P. bryhnii (Steph.) Inoue, P. oppositus (Reinw., Blume et Nees) S. Hatt., P. fimbriatus (Mitt.) Inoue
Subclass Marchantiidae (ゼニゴケ亜綱)
450 species and 27 genera in 12 families in the world (39 species and 18 genera in nine families in Japan)
Conocephalaceae (ジャゴケ)
Conocephalum conicum (L.) Dum. (ジャゴケ), 1属1種
Conocephalum Conocephalum Conocephalum
Fig. C. conicum at the backyard of GSEES, HU. Left/Center, on May 1 2021. Right, on April 22 2022
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.
M. papillata Raddi. ssp. grossibarba (Steph.) Bischl. (Syn. M. tosana Steph.): This species had been reported from the west of Kanto District, and was recorded from Mt. Usu, Hokkaido, northern Japan. The subspecies grows on soil around fumaroles in the mountain. (樋口・露崎 2023)
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

Ecology (生態)

Comparative cryptogam ecology, in particular bryophyte and lichen (Cornelissen et al. 2007)
  1. contribute substantially to above-ground biomass (lichens, bryophytes)
  2. host nitrogen-fixing bacteria, providing major soil N input (lichens, bryophytes)
  3. control soil chemistry and nutrition through the accumulation of recalcitrant polyphenols (bryophytes) and through their control over soil and vegetation hydrology and temperatures
  4. both promote erosion (rock weathering by lichens) and prevent it (biological crusts in deserts)
  5. provide a staple food to mammals such as reindeer (lichens) and arthropodes, with important feedbacks to soils and biota
  6. both facilitate and compete with vascular plants

[Species checklist in Alaska]


Moss flora list on Poker Flat (after the 2004 wildfire)
The specimens are stored in SAPS. Thank Dr. Higuhi for the identification!
  • 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. (スギバミズゴケ)
  • 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]