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Pteridophyta (羊歯植物)

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

Division Pteridophyta = ferns, sensu lato
The records of number of fern species
Linne's Era: 11 gen. 190 spp.
1793 Smith JE: 13 new genera → total = 24 genera

Presence or absence of annulus (環帯)

1806 Schwarz: "Synopsis Filicum" → 36 gen. 700 spp
1810 Willdenew: "Linne's Species Plantarum (5 edn)" → 41 gen. 1010 spp. 1961 Cronquist A:

Subkingdom II Embryophyta Division 9 Bryophyta
Division 10 Psilophyta (マツバラン) __ Class 10a Psilotae
Division 11 Lepidophyta (ヒカゲノカズラ) __ Class 11a Lycopodiae
Division 12 Colamophyta (トクサ) __ Class 12a Equisetae
Division 13 Filicophyta (シダ) __ Class 13a Filices

Life cycle
Rreproductive organ = sporangium(-gia) → spore
Spory (胞子型)
a) Homosporous type (homospory)
life cycle
Ex. Pteridium aquilinum, Athyrium niponicum: divide primodal cell into two parts
life cycle b) Heterosporous type (heterospory)
microsporanium → microspore / macrosporanium → macrospore

┌─→ n (female gametophyte) → egg (n) ─────┐
│┌→ n (male gametophyte) ─→ spermatozoid (n)─┤
││     n ←─┐                             2n sporophyte
│└── n ←─┼─ microsporangium (2n) ←┐      │
│       n ←─┘                          │      ↓
│   microspore                            ├ 2n sporophyte
│       n ←─┐                          │
└─── n ←─┼─ macrosporangium (2n) ←┘
         n ←─┘

⇒ Engler's syllabus (エングラー体系), Gymnospermae, Angiospermae

moss cycle cp. Life cycle of moss
Pteridophyte coefficient (シダ係数)
= pteridophyte index

related to climate (high air moisture and/or temperature)

= (25 × number of fern species)/(number of seed plant species)

[ morphology of seed plants]

Morphology (形態)

External morphology (外部形態)

Morphology Morphology
Frond (葉状体) ≈ leaf (葉) for fern, cydad and palm: The leaf of the fern bearing the pinnae

= blade + stipe

Blade (葉身): part of frond bearing the foliage
Stipe (柄/茎) or petiole (葉柄): leaf stalk of the fern arising from the rhizome and below the blade
Apex (頂端): the top of blade
Midrib (中肋): central or principal vein of a frond

Number of veins = the number of midrib branches of the fronds

Pinnule(s) (小羽片) ≈ leaflet for seed plants
Pinna (pl. -e) (羽片): the primary division of a compound fern frond. A distinct subdivision of the blade, when the blade segments are fully divided to the rachis
Rachis (羽軸): the main stalk or central vein, of a fern frond that bears the pinnae - the blade section. The section below the pinnae is called the stipe
Sorus (pl. sori) (胞子嚢群): a cluster of sporangia arranged on the underside or margins of the pinnae in a distinctive pattern in each fern genus
Indusium (pl. -a) (包膜): membranes covering the sori, forming a net-like or skirt-like shape
Sporangium (pl. -a) (胞子嚢) for mosses and ferns: a spore case in which the spores develop. Thin walled, usually stalked
Spore (胞子): a minute, typically one-celled, reproductive unit capable of giving rise to a new individual without sexual fusion

Internal morphology (内部形態)

Stele (中心柱)

stele Protostele    Siphonostele    Solenostele    Dicyostele
Fig. Types of vascular cylinder (stele) in ferns. Black dots: phloem. Orange, meshed areas: xylem

[*: extinction]

Division Pteridophyta (シダ植物門)

Phylogeny constructed by PPG I (2016)
Rhyniopsida* リニア
Trimerophytopsida* トリメロフイトン
Zosterophyllopsida* ゾステロフィルム
Psilotopsida マツバラン

Psilotales マツバラン

Lycopsida ヒカゲノカズラ
Drepanophycales* ドレパノフイクス
Protolepidodendrales* 古生リンボク目(プロトレピデンドロン目)
Lepidodendrales* リンボク(トレピデンドロン)
Lycopodiales ヒカゲノカズラ
Selaginellales イワヒバ
Pleuromeiales プレウロメイア
Isoetales ミズニラ
Equisetopsida トクサ綱
Hyeniales* ヒエニア
Pseudoborniales* プセウドボルニア
Sphenophyllales* スフェノフィルム
Equisetales トクサ目
Equisetaceae (トクサ)
Filicopsida シダ綱
Cladoxylales* クラドキシロン
Iridopteridales* イリドプテリス
Rhacophytales* ラコフイトン
Stauropteridales* スタウロプテリス
Zygopteridales* ジゴプテリス
Marattiales リュウビンタイ
Ophioglossales ハナヤスリ
Ophioglossaceae ハナヤスリ
Filicales シダ
Progymnospermopsida* 前裸子植物
Aneurophytales* アネウロフイトン
Archaeopteridales* アルカエオプテリス
Protopityales* プロトピチス

Subdivision Psilophytina (裸茎植物亜門), or Division Psilophyta

Dichotomy (叉状分枝)
Protostele (原生中心柱)
Terminal sporangium

Class Psilophytopsida (古生マツバラン綱)*


Class Psilotopsida (マツバラン綱)

The phylogenic relationship to Psilophytopsida is negative
Exoscopic embryo
Leafless and rootless
Protostele Psilotum
1-5: Psilotum triquetrum, 1 oberer Tell der Pflanze mit Sporangien, 2 geschlossenes Sporangium, 3 aufgesprung enes Sporangium, 4 Sporen, 5 Querschnitt durch ein noch unreifes Sporangium, 6 Spros, 6-7 Tmesipteris Sporophyll
Order Psilotales (マツバラン)
degeneration of microphyll → considered to be in microphyllous fern Psilotaceae (マツバラン)
Psilotum L.
P. nudum (L.) P. Beauv. (マツバラン)
P. complanatum Sw. (ソウメングサ)
Fig. Psiltoum triquetrum (Gametophyte)
Tmesipteridaceae (イヌナンカクラン), moved into Psilotaceae
Tmesipteris Berhn., hanging fork fern

Subdivision Lepidophytina (小葉植物亜門)

= Division Lepidophyta (Lycopodophyta) (小葉植物植物門), clubmosses and quillworts
400-600 spp. in the world, 17 spp. in Japan, and 11 spp. in Hokkaido
Spore: germination, growth

Gametophyta, e.g., Licopodium complanatum
Licopodium complanatum

palisade tissue (柵状組織)
storage tissue (貯蔵組織)

The development of zygote

suspensor (胚柄): non-divisive

Class Lepidopsida (小葉植物綱)

⇒ Aglossopsida (無舌綱) when Glossopsida is established
Order Protolepidodendrales (古生ヒカゲノカズラ*)
Order Lycopodiales (ヒカゲノカズラ)
Carboniferous - present
Lycopodium L. (ヒカゲノカズラ)

Class Glossopsida (有舌綱)

Order Lepidodendrales (リンボク目)*
Devonian - Permian
Order Lepidocarpales
This order is discarded, when Lepidocarpaceae → Lepidodendales, Maidesmiaceae → Selaginellales
Lepidocarpaceae*: Ledpidocarpon Scott
Maidesmiaceae*: Miadesmia Betr.
Order Isoetales ミズニラ
Reproductive organs developed on the base of leaves in autumn (microphyllous) ↔ leaves produced in spring are vegetative organs only
heterospory (異型胞子): macrospore – developing ealier, microspore – later
eusporangiate (真嚢): germination Isoetales

macrospore → free nucloeous division
microspore → protharial cell


Pleuromeia (Triassic): dioecious - related to Selaginellales (Mägdefran 1931, 43)
Nathorstiana (Cretaceous)
Stylites: heterosproy, fern-typed sperm (Rauh 1954)
Isoetaceae (ミズニラ)
Isoetes (ミズニラ): I. australis S.Williams (WA), I. drummondii A. Braun (WA), I. tripus A. Braun (WA)
Stylites: discovered on the mountain of Pelu in 1954

Microphyllous → spiral microphyllous group = Isoetes / Stylites

Order Selaginellales (イワヒバ)
Carboniferous - present
Subterraniean gametophyte: mycotrophy (菌栄養) not developing chlorophyll
Soprangium developing on the upper surface of sporophyll → phylogenetically different from Lycopodium
Differentiating leaves, stems and roots → Protostele
Stem: slim = exarch (外原型) ↔ thick = endarch (内原型) Cf. Botryopteris (古生シダ)
Root: rhizophore (担根体) by exogenous branching (or growth)

← roots are generally formed by endogenous branching
Williams (1933-38): forming stem or rhizophore is determined by the amount of hormone (auxin)
Webster & Steeve (1963, 1964, 1967): root cap is detached after the formation of root cap in the early developmental stages (no replication study)

Microphyllous leaf (小葉)

ligule (小舌): function unknown, absent in Lycopodiales
ventral leaf (腹葉): large leaves, opened to both the sides, of the leaves arranged on four liens
dorsal leaf (背葉): small leaves, arranged on two lines, coherent at dorsal side

Selaginella P. Beauv. (イワヒバ) in Japan

S. biformis A. Br. ex Kuhn (ツルカタヒバ)
S. boninensis Baker (ヒバゴケ)
S. doederleinii Hieron. (オニクラマゴケ)

var. opaca Seriz. (コウズシマクラマゴケ)

S. helvetica (L.) Link (エゾノヒメクラマゴケ)
S. heterostachys Baker (ヒメクラマゴケ)
S. involvens(Sw.) Spring (カタヒバ), syn. S. gracillima (Kunze) Alston (Western Australia)
S. leptophylla Baker (コケカタヒバ)
S. limbata Alston (アマミクラマゴケ)
S. lutchuensis Koidz. (ヒメムカデクラマゴケ)
S. moellendorffii Hieron. (イヌカタヒバ)
S. nipponica Franch. et Savat. (タチクラマゴケ)
S. remotifolia Spring (クラマゴケ)
S. selaginoides (L.) Link (コケスギラン)
S. shakotanensis (Franch. ex Takeda) Miyabe et Kudo (ヒモカズラ)
S. sibirica (Milde) Hieron. (エゾノヒモカズラ)
S. tamariscina (Beauv.) Spring (イワヒバ), south to Oshima Peninsula and Okushiri Island
S. tama-montana Serizawa (ヤマクラマゴケ)
S. uncinata (Desv.) Spring (コンテリクラマゴケ)

Subdivision Sphenophytina (有節植物亜門), or Division Arthrophyta (Calamophyta)

all of the present species are microphyllous type
homospory (morophological): no suspensor, embryo developing outward

Sporophyll (= sporangiophore)
♂ gametophyte, ♀ gametophyte: dioecious

Class Arthropsida (有節植物綱)

horsetail and sconring rushes
Order Protoarticulales (Hyeniales)*
Hyeniaceae*: Devonian: Hyenia (mid-Devonian)
Calamophytaceae: identified by brancing patterns and separated from Equisetaceae

formerly in the same group with Equisetaceae

Order Sphenophyllales*
Devonian -Triassic
Spehno- [wedge] + phyllum [leaf] → sphenophyllous plant – herb-like

Protostele → siphonostele (管状中心柱)
Leaf: verticillate, dimorphic – telome origin (macrophyllous)
Equisetum________Sphenophyllum ⇒ dimorphic leaves

Order Cheirostrobales*
Order Pseudoborniales*
Order Calamitales (ロボク)*
Devonian - Triassic
Secondary auxetic growth - remarkable → stem: 30 cm φ, 20-30 m long
Calamitaceae ロボク
leaves primarily authorized as Annularia → later, combined with stem of which name is Calamites
Calamites Suckow (ロボク): remarkable macrophyllous leaf
Order Equisetales (トクサ)
Carboniferous - present
Equisetaceae (トクサ)
Equisetum L. (トクサ): only this genus is present now
Chromosome number: n = 108
Herb-like = no secondary auxetic growth
Tarminal sporangia = marginal
Troll (1928), Göbel (1930)

Strobus on Equisetum is "one" flower

Scott (1912), Eames (1936), Ogura (1937)

Counter by using Calamostachys and Paleostachya

Equisetum L., horsetail (トクサ)

E. arvense L. (スギナ)
E. fluviatile L. (Syn. E. limosum L.) (ミズドクサ)
E. hyemale L. (トクサ)
E. palustre L. (イヌスギナ)
E. pratense Ehrh. (ヤチスギナ)
E. ramosissimum Desf. (イヌドクサ)

var. ripense (Nakai et F. Maekawa) Tagawa (タカトクサ)

E. scirpoides Michx. (ヒメドクサ)
E. sylvaticum L. (フサスギナ)
E. variegatum Schleich. ex Weber et Mohr (チシマヒメドクサ)
Putative hybrids
E. × moorei Newn. = E. ramosissimum × hyemale (トリトクサ)
E. × rothmaleri C. N. Page = E. palustre × arvense (フォーリースギナ)
E. × litorale Kuhlew. ex Rupr. = E. fluviatile × arvense (ハマスギナ)

Class Filicopsida (真正シダ綱)


Fig. Ontogeny and structure of the two principal types of sporangia in vascular plants. a-e, the eusporangium; a'-g', the leptosporangium.

Subclass Primofilicidae (原シダ亜綱*)

from mid-Devonian, flourish in Carboniferous

Subclass Filiciedae 真正シダ亜綱

a) Eusporangiate group (真嚢シダ類)
= Ophioglossales + Marattiales

Ex. Ophioglossum, Botrychium
One leaf, complex veins____________Several leaves, branching

b) Leptosporangiate (薄嚢シダ類)
= Filicales (真正シダ目) + Marsileales + Salviniales
Fig. Life cycle of a fern, Polypodium vulgare (オオエゾデンダ) (Engler's Syllabas 1954)
Order Eusporangiales (真嚢シダ目)
Ophioglossaceae (ハナヤスリ): 4 gen. 70 spp. in the world
Botrychium Sw. (ハナワラビ)
Ophioglossum L. (ハナヤスリ): O. lusitanicum L. (WA)
Schizaeaceae Kaulf. (フサシダ)
Schizaea Smith (S. fistulosa Labill.)
Lygodiaceae C. Presl (カニクサ), when separated from Schizaeaceae
Lygodium Sw. (カニクサ, climbing fern), one genus when this family is established: native to the tropical regions, consisting of ca 40 species

Japan: L. circinnatum (Burm. f.) Sw. (ナンゴクカニクサ). L. flexuosum (L.) Sw. (シマカニクサ). L. japonicum (Thunb.) Sw. (カニクサ). L. microphyllum (Cav.) R. Br. (イリオモテシャミセンヅル), Yaeyama Islands

Marattiaceae (リュウビンタイ)

Angiopteris Hoffm. リュウビンタイ (A. lygodiifolia リュウビンタイ)

Psaroniaceae (fossil species): Psaronius Cotta
Order Protolepidosporangiales 前薄嚢シダ
+ Osumudales ゼンマイ (included or separated)

Eusporangiagae → Seed plant (真嚢)

Osmundaceae (ゼンマイ): Permian-present
Macrophyllous → morphologically-close to leptosproangiatae
Stalk on sporangium becoming thick → sori developed only on sporophylls because of the specialization of sorus morphology
Tapetum (絨毯細胞) revmoed and the outer layer remained → two-layered outer wall
Homosporic: 300-500 spores

Osmunda L. (ゼンマイ)

Order Leptosporangiales (薄嚢シダ)
1954 German: 14 family, 5000-7000 spp.
Loxoma R. Br. ex A. Cunn., Loxomopsis Christ
Pteridaceae Kirchn. (イノモトソウ)
Sori on the leaf margin
Forming coenosorus (thickened, roll-up inside)
Rhizome = siphonostele
Onychium Kaulfuss. (タチシノブ), sometimes separated from Pteridaceae by establishing Parkeriaceae Hook. (ホウライシダ) that is not allowed to use due to the rule of ICBN
one species in Japan (10 species in the world)
O. japonicum (Thunb.) Kunze (タチシノブ)

Fig. Dennstaedtia wilfordii
(Kurata & Nakaike 2005)

Dennstaedtiaceae (コバノイシカグマ, separated from Pteridaceae) Parkeriaceae or Adiantaceae (ホウライシダ)
Adiantum L. (クジャクシダ)
Anogramma Link: A. leptophylla (L.) Link, Western Australia
Cheilanthes Sw., mostly in Western Australia: C. distans (R. Br.) Mett., C. sieberi Kunze, C. austrotenuifolia H. Quirk et T. C. Chambers
Hymenophyllopsidaceae (コケシノブ)
Hymenophyllum Smith (コウヤコケシノブ): H. barbatum (コウヤコケシノブ)
Davalliaceae (シノブ)
Plagiogyriaceae (キジノオシダ)
Plagiogyria (Kunze) Mett. (キジノオシダ) (one genus in this family)
Cyatheaceae (ヘゴ), merged into Aspidiaceae, s.l.)
Sporocarp, sorus: Marsilea-typed emergence (as well as Dicksoniaceae タカワラビ)

Cyathea Smith ヘゴ: C. fauriei ヘゴ、tree-like, evergreen
Sphaeropteris Bernth.: S. cooperi (Hook. et F. Muell.) R. Tyron (Western Australia)

Thelypteridaceae (ヒメシダ), Western Australia

Cyclosorus Link
Thelypteris Schmidel

Dryopteridaceae (s.s.), Aspidiaceae (s.l.) (オシダ)
Arachniodes Blume (カナワラビ), holly fern (syn. Polystichopsis (J. Sm.) Holtt., カナワラビ)
Dryopteris Adans. (オシダ)
Polystichum Roth (イノデ): 200 spp. in world, 23 spp. in Japan
Matteuccia Tadaro (クサソテツ)
Onoclea L. (コウヤワラビ): two species in Japan

Fig. Diplazium sibiricum
var. glabrum
Diplazium squamigerum
(Kurata & Nakaike 2005)

Oleandraceae ツルシダ (separated from Dryopteridaceae, or merged into Davalliaceae)

Nephrolepis cordifolia (L.) Presl (タマシダ)

Blechoraceae (シシガシラ)
Lindsaeaceae: Lindsaea Dryander ex Smith (L. linearis Sw., Western Australia)
Aspleniaceae (チャセンシダ): > 650 spp. in 2 genera

Asplenium L. (チャセンシダ): A. aethiopicum (Burm. f.) Bech., Western Australia
Pleurosorus Fee: P. rutifolius (R. Br.) Fee, Western Australia

Gleicheniaceae (ウラジロ), included into Aspidiaceae, s.l.
Woodsiaceae イダデンダ, Athyrium is included in this family when established
Athyriaceae (メシダ), included in Aspidiaceae, s.l. when established
Matoniaceae (マトニア)

Matonia R. Br.
Phanerosorus Copel.

Polypodiaceae (ウラボシ)
Vittariaceae (シシラン)
Order Hydropteridales
Cretaceous or Triassic-present (sometimes, dividied into two orders, Marsileales/Salviniales)
Common characteristics: sporocarp = sorus enveloped by indusium
Different characteristics: Salviniales = float, annuals ⇔ Marsileales = submerged, perennials
Order Marsileales (デンジソウ) when established
Marsileaceae (デンジソウ)

Marsilea L. デンジソウ → quadrifolia

Order Salviniales (サンショウモ), when established
Salviniaceae (サンショウモ)
Azollaceaee (アカウキクサ)

Azolla Lam. (アカウキクサ) (A. caroliniana, A. imbricata (アカウキクサ), A. pinnata)

Vertical section_________________Cross section (sporocarp)

3rd species = heterosporic, spore formation and germination morphology are the same among the three species


Genus (属)

Botrychium Sw. (ハナワラビ) in Japan

B. atrovirens (Sahashi) M. Kato (シチトウハナワラビ)
B. boreale (Fr.) Milde (タカネハナワラビ)
B. daucifolium Wall. ex Hook. et Grev. (シマオオハナワラビ)
B. formosanum Tagawa (ホウライハナワラビ)
B. japonicum (Prantl) Underw. (オオハナワラビ)
B. lanceolatum (S. G. Gmel.) Angstr. (ミヤマハナワラビ)

ssp. angustisegmentum (Pease et A. H. Moore) Clausen (ホソバミヤマハナワラビ)

B. lanuginosum Wall. ex Hook. et Grev. (アリサンハナワラビ)
B. lunaria (L.) Sw. (ヒメハナワラビ)
B. microphyllum (Sahashi) K. Iwats. (イブリハナワラビ)
B. multifidum (S. G. Gmel.) Rupr. (ヤマハナワラビ)
B. nipponicum Makino (アカハナワラビ)

var. minus (H.Hara) K. Iwats. (ウスイハナワラビ)

B. simplex E. Hitchc. var. tenebrosum (A. A. Eaton) Clausen (コケハナワラビ)
B. strictum Underw. (ナガホノナツノハナワラビ)
B. ternatum (Thunb.) Sw. (フユノハナワラビ)
B. triangularifolium (Sahashi) M. Kato (ミドリハナワラビ)
B. virginianum (L.) Sw. (ナツノハナワラビ)
B. japonicum (Prantl) Underw. × B. ternatum (Thunb.) Lyon (アイフユハナワラビ)
B. × argutum (Sahashi) K.Iwats., nom. nud. (ハブハナワラビ)
B. × elegans (Sahashi) K.Iwats., nom. nud. (アカネハナワラビ)
B. × longistipitatum (Sahashi) K.Iwats., nom. nud. (アイイズハナワラビ)
B. × pulchrum (Sahashi) K.Iwats., nom. nud. (アンコハナワラビ)

Ophioglossum L. (ハナヤスリ)

O. austroasiaticum M. Nishida (タイワンハナヤスリ)
O. kawamurae Tagawa (サクラジマハナヤスリ)
O. namegatae M. Nishida et Kurita (トネハナヤスリ)
O. parvifolium Grev. et Hook. (イオウジマハナヤスリ)
O. parvum M. Nishida et Kurita (チャボハナヤスリ)
O. pendulum L. (コブラン)
O. petiolatum Hook. (コヒロハハナヤスリ)
O. thermale Kom. (ハマハナヤスリ)
O. vulgatum L. (ヒロハハナヤスリ)
O. × nipponicum Miyabe et Kudô (コハナヤスリ)

Osmunda L. (ゼンマイ) in Japan

O. banksiifolia (Pr.) Kuhn (シロヤマゼンマイ)
O. cinnamomea L. (ヤマドリゼンマイ)
O. claytoniana L. (オニゼンマイ)

var. vestita (Wall.) Milde (タイワンオニゼンマイ)

O. japonica Thunb. (ゼンマイ)
O. lancea Thunb. (ヤシャゼンマイ): endemic to Japan, including southern Hokkaido
O. nipponica Makino (コゼンマイ)

var. divisa (Makino) Nakai (シシゼンマイ)

O. × intermedia (Honda) Sugimoto (オオバヤシャゼンマイ)

Metzgar JS, Skog JE, Zimmer EA & Pryer KM. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany 33: 31-36