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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
life cycle

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

moss cycle cp. Life cycle of moss

*: extinction ⇒ geographical timescale

Division Pteridophyta (シダ植物門)

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
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
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.
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: 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

Equisetaceae (トクサ)
Equisetum L. (トクサ)

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 (ハナヤスリ), Botrychium (ハナワラビ)
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)
Dennstaedtiaceae (コバノイシカグマ, separated from Pteridaceae)

Fig. Dennstaedtia wilfordii
(Kurata & Nakaike 2005)

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)
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)
Dryopteris Adans. (オシダ)
Polystichum Roth (イノデ): 200 spp. in world, 23 spp. in Japan
Polystichopsis (J. Sm.) Holtt.(カナワラビ)
Matteuccia Tadaro (クサソテツ)
Onoclea L. (コウヤワラビ): two species in Japan
O. orientalis (Hook.) Hook. (イヌガンソク)
O. sensibilis L. (コウヤワラビ, s.l.)

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

Cystopteris (ナヨシダ): C. sudetica A. Br. et Milde (ヤマヒメワラビ), discovered in Engaru (遠軽) by H Hayashi (林 廣志), C. fragilis (L.) Bernh. ナヨシダ
Diplazium Swartz (ヘラシダ): D. sibiricum (Turcz. ex Kunze) Kurata キタノミヤマシダ (var. glabrum (Tagawa) Kurata ミヤマシダ, long rhizomes)

Athyriaceae (メシダ), included in Aspidiaceae, s.l. when established

Athyrium Roth (メシダ): A. pycnosorum Christ ミヤマシケシダ

Matoniaceae (マトニア)

Matonia R. Br.
Phanerosorus Copel.

Polypodiaceae (ウラボシ)
Lemmaphyllum Presl. (マメヅタ)

L. microphyllum C. Presl (マメヅタ), L. nobukoanum (Makino) Ching (ヒメマメヅタ), recorded from Taiwan, L. pyriforme (Ching) Ching (オニマメヅタ)

Pyrrosia Mirbel (ヒトツバ)
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

Indusium (包膜): membranes, forming a net-like or skirt-like shape, covering the sori

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)
Salviniaceae (サンショウモ)

Salvinia Adans. (S. natans 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)

Athyrium Roth (メシダ)

Approximately 180 species in the world
A. alpestre (Hoppe) Clairv. (オクヤマワラビ)
A. arisanense (Hayata) Tagawa (タイワンアリサンイヌワラビ)
A. atkinsonii Bedd. (テバコワラビ)
A. brevifrons Nakai ex Tagawa (エゾメシダ)
A. clivicola Tagawa (カラクサイヌワラビ)
A. delavayi Christ (ホウライイヌワラビ)
A. deltoidofrons Makino (サトメシダ)

f. acutissimum (Kodama) Sa. Kurata (トガリバメシダ)
f. ohmurae Sa. Kurata (オゼサトメシダ) A. deltoidofrons Makino × iseanum Rosenst. (ホソバサトメシダ)
A. deltoidofrons Makino × otophorum (Miq.) Koidz. (タニサトメシダ)
A. deltoidofrons Makino × wardii (Hook.) Makino (ギフイヌワラビ)

A. epirachis (Christ) Ching (ヘイケイヌワラビ)
A. filix-femina (L.) Roth (セイヨウメシダ), the type
A. frangulum Tagawa (ミヤコイヌワラビ)
A. iseanum Rosenst. (ホソバイヌワラビ), south to Akita and Ibaragi Prefecture
A. kenzo-satakei Sa. Kurata (シビイヌワラビ)
A. kirisimaense Tagawa (キリシマヘビノネゴザ)
A. kuratae Seriz. (ツクシイヌワラビ)
A. melanolepis (Franch. et Sav.) Christ (ミヤマメシダ)
A. nakanoi Makino (ヒメホウビシダ)
A. neglectum Seriz. (コシノサトメシダ)
A. nikkoense Makino (イワイヌワラビ)
A. niponicum (Mett.) Hance (イヌワラビ)
A. otophorum (Miq.) Koidz. (タニイヌワラビ)
A. oblitescens Sa. Kurata (サキモリイヌワラビ)
A. palustre Seriz. (サカバサトメシダ)
A. pinetorum Tagawa (タカネサトメシダ)
A. reflexipinnum Hayata (サカバイヌワラビ)
A. rupestre Kodama (ミヤマヘビノネゴザ)
A. strigillosum (Lowe) T. Moore ex Salomon (コモチイヌワラビ)
A. spinulosum (Maxim.) Milde (カラフトミヤマシダ)
A. sheareri (Baker) Ching (ウラボシノコギリシダ)
A. tozanense (Hayata) Hayata (シマイヌワラビ)
A. silvicola Tagawa (タカサゴイヌワラビ)
A. vidalii (Franch. et Sav.) Nakai (ヤマイヌワラビ)
A. viridescentipes Sa. Kurata (アオグキイヌワラビ)
A. wardii (Hook.) Makino (ヒロハイヌワラビ), south to Ibaragi and Toyama Prefecture

var. inadae Tagawa (ルリデライヌワラビ)
f. chloropodum Sa. Kurata (ミドリヒロハイヌワラビ)

A. yakusimense Tagawa (ヤクシマタニイヌワラビ)
A. yokoscense (Franch. et Sav.) Christ (ヘビノネゴザ)

summergreen. roky sites - forest floor (diverse habitats, e.g., mines. Central-eastern China - Manchuria - Korea - Amur - Sakhaline - southern Kuril - Japan

A. × multifidum Rosenst. (オオサトメシダ)

Goniophlebium (Blume) C. Presl (ナンヨウウラボシ, tentative)

G. amamianum (Tagawa) Nakaike (アマミアオネカズラ)
G. amoenum (Wall. ex Mett.) Bedd. (アリサンデンダ)
G. formosanum (Baker) Rödl.-Linder (タイワンアオネカズラ)
G. mengtzeense (Christ) Rödl-Linder (タイワンウラボシ)
G. microrhizoma (C. B. Clarke ex Baker) Bedd. (アリサンウラボシ)
G. niponicum (Mett.) Bedd. (アオネカズラ), syn. Polypodium nipponicum Mett.
G. someyae (Yatabe) Ebihara (ミョウギシダ)

var. awaense (Tagawa) Nakaike (アワミョウギシダ)

Southeastern Asia
G. verrucosum (Wall. ex Hook.) J. Sm. (ナンヨウウラボシ, tentative translation), unresolved name

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

Dryopteris Adans. (オシダ)

250 spp mostly in the temperate Northern Hemisphere
Characterized by a vase-like ring of fronds
D. austriaca (Jacq.) Woynar ex Schinz. et Thell. (シラネワラビ)
D. bissetiana (Baker) C. Chr. (ヤマイタチシダ)
D. crassirhizoma Nakai (オシダ)
D. erythrosora (D. C. Eaton) Kuntze (ベニシダ)
D. filix-mas (L.) Scott (セイヨウオシダ, male fern), not in Japan
D. lacera (Thunb.) Kuntze (クマワラビ)
D. monticola (Makino) C. Chr. (ミヤマベニシダ)
D. nipponensis Koidz. (トウゴクシダ)
D. sabae Koidz. (ミヤマイタチシダ)
D. tokyoensis (Makino) C. Chr. (タニヘゴ)
D. varia (L.) Kuntze (ナンカイイタチシダ)
D. × kominatoensis Tagawa (タニヘゴモドキ)

A putative hybrid between D. tokyoensis and D. monticola

Thelypteris Schmidel (ヒメシダ)

T. acuminata (Houtt.) C. V. Morton (ホシダ)
T. adscendens Ching (ヒカゲヤワラシダ)
T. angulariloba Ching (オオハシゴシダ)
T. angustifrons (Miq.) Ching (コハシゴシダ)
T. arida (D. Don) C. V. Morton (タイワンホシダ)
T. auriculata (J. Sm.) K. Iwats. (オオミゾシダ)
T. aurita (Hook.) Ching (ハイミミガタシダ)
T. beddomei (Baker) Ching (ホソバショリマ)
T. boninensis (Kodama ex Koidz.) K. Iwats. (オオホシダ)
T. bukoensis (Tagawa) Ching (タチヒメワラビ)
T. castanea (Tagawa) Ching (タイワンハシゴシダ)
T. cystopteroides (D. C. Eaton) Ching (ヒメハシゴシダ)
T. decursivepinnata (H. C. Hall) Ching (ゲジゲジシダ)
T. dentata (Forssk.) E. P. St. John (イヌケホシダ)
T. dictyoclinoides (Ching) C. F. Reed (アミシダモドキ)
T. erubescens (Wall. ex Hook.) Ching (タイヨウシダ)
T. esquirolii (Christ) Ching

var. esquirolii (オオイブキシダ)
var. glabrata (Christ) K. Iwats. (イブキシダ)

T. flexilis (Christ) Ching (トサノミゾシダモドキ)
T. glanduligera (Kunze) Ching (ハシゴシダ)
T. gracilescens (Blume) Ching (シマヤワラシダ)
T. griffithii (Hook. f. et Thomson) C. F. Reed

var. griffithii (オオアミシダ)
var. wilfordii (Hook.) C. M. Kuo (アミシダ)

T. gymnocarpa (Copel.) C. V. Morton

ssp. amabilis (Tagawa) C. V. Morton (ヒメミゾシダ)

T. gymnopteridifrons (Hayata) C. M. Kuo (トネリコシダ)
T. hattorii (H. Ito) Tagawa (ヨコグラヒメワラビ)

var. nemoralis (Ching) Sa. Kurata (ツクシヤワラシダ)

T. interrupta (Willd.) K. Iwats. (テツホシダ)
T. jaculosa (C. Chr.) Panigrahi (クシノハシダ)
T. japonica (Baker) Ching (ハリガネワラビ)
T. latipinna (Benth.) K. Iwats. (ヒロハホシダ)
T. laxa (Franch. et Sav.) Ching (ヤワラシダ)

f. glabrescens H. Ito (ウスゲヤワラシダ)

T. leucolepis (C. Presl) Ching (オニヒメワラビ)
T. leveillei (Christ) C. M. Kuo (ミゾシダモドキ)
T. liukiuensis (Christ ex Matsum.) K. Iwats. (オオコウモリシダ)
T. longipetiolata (K. Iwats.) K. Iwats. (ナガエコウモリシダ)
T. miyagii (H. Ito) Nakato, Sahashi et M. Kato (リュウキュウハシゴシダ)
T. musashiensis (Hiyama) Nakato, Sahashi et M. Kato (イワハリガネワラビ)
T. nipponica (Franch. et Sav.) Ching (ニッコウシダ)
T. ogasawarensis (Nakai) H. Ito ex Honda (ムニンヒメワラビ)
T. oligophlebia (Baker) Ching (トウヒメワラビ)
T. palustris (Salisb.) Schott (ヒメシダ)
T. papilio (Hope) K. Iwats. (ウスバクシノハシダ)
T. parasitica (L.) Tardieu (ケホシダ)
T. phegopteris (L.) Sloss. ex Rydb. (ミヤマワラビ)
T. pozoi (Lag.) C. V. Morton

ssp. mollissima (Fisch. ex Kunze) C. V. Morton (ミゾシダ)

T. productus (Kaulf.) C. F. Reed (コウトウシダ)
T. prolifera (Retz.) C. F. Reed (ツルアミシダ)
T. pseudoacuminata Shimura, nom. nud. (ホシダモドキ)
T. pyrrhorhachis (Kunze) B. K. Nayar et Kaur (ニイタカワラビ)
T. quelpaertensis (Christ) Ching (オオバショリマ)

var. yakumontana (Masam.) Tagawa (ヤクシマショリマ)

T. simplex (Hook.) K. Iwats. (ヒトツバコウモリシダ)
T. subaurita (Tagawa) Ching (ミミガタシダ)
T. taiwanensis (C. Chr.) K. Iwats. (コバザケシダ)
T. torresiana (Gaudich.) Alston

var. torresiana (アラゲヒメワラビ)

T. triphylla (Sw.) K. Iwats. (コウモリシダ)

var. parishii (Bedd.) K. Iwats. (ホソバコウモリシダ)

T. truncata (Poir.) K. Iwats. (ナタギリシダ)
T. uraiensis (Rosenst.) Ching (タイワンハリガネワラビ)
T. viridifrons Tagawa (ミドリヒメワラビ)
T. acuminata (Houtt.) C. V. Morton × T. parasitica (L.) Tardieu (アイノコホシダ)
T. japonica (Baker) Ching × T. musashiensis (Hiyama) Nakato, Sahashi et M. Kato (アイハリガネワラビ)
T. × incesta W. H. Wagner (カワバタホシダ)
T. × insularis (K. Iwats.) K. Iwats. (エラブコウモリシダ)
T. × pseudoliukiuensis Seriz. (ヤエヤマコウモリシダ)
T. × subviridifrons Seriz. (アイヒメワラビ)