Polypodiales

Classification

└Tracheophytes

└Euphyllophytes

└Polypodiophyta

└Ferns (Polypodiopsida)

└Polypodiidae

└Polypodiales

Geologic Range

• Cretaceous - present

Dennstaedtiaceae

• 10 extant genera (Blotiella, Coptodipteris, Dennstaedtia, Histiopteris, Hypolepis, Leptolepia, Microlepia, Monachosorum, Oenotrichia, Paesia, and Pteridium) with ~240 known species

• Ferns are found worldwide, but mostly pantropical, except Pteridium and Dennstaedtia

• Cretaceous - present

Stems

• Rhizomatous

Leaves

• Fronds are large and highly-divided

Reproductive features

• Leaves have either small, round intramarginal sori with cup-shaped indusia (e.g. Dennstaedtia) or linear marginal sori with a false indusium formed from the reflexed leaf margin (e.g. Pteridium)

Lindsaeaceae

• 6-7 extant genera (Lindsaea, Nesolindsaea, Odontosoria, Osmolindsaea, Sphenomeris, Tapeinidium, and Xyropteris); ~220 species

• Ferns are mostly pantropical, but also extend into temperate regions of eastern Asia, New Zealand, and South America

• Cretaceous - present

Stems

• Rhizomes short to long creeping with nonclathrate scales or uniseriate hairs

Leaves

• Fronds 1-3x pinnate or more divided; veins usually free

Reproductive features

• Sori marginal or submarginal

• Indusia open towards the margin, sometimes attached at sides, or sori covered by the reflexed segment margin

Lonchitidaceae

• 1 genus (Lonchitis); 30 species

• Ferns with neotropical distribution

Stems

• ?

Leaves

• ?

Reproductive features

• ?

Pteridaceae

• 5 subfamilies; 45 genera; 1,150 species

• Mostly terrestrial or epipetric (growing on rock).

Cheilanthoideae

• Adiantopsis, Aleuritopteris, Allosorus, Argyrochosma, Aspidotis, Astrolepis, Bommeria, Calciphilopteris, Cheilanthes, Cheiloplecton, Doryopteris, Gaga, Hemionitis, Lytoneuron, Mildella, Myriopteris, Notholaena, Ormopteris, Paragymnopteris, Parahemionitis, Pellaea, Pentagramma, and Trachypteris

Cryptogrammoideae

• Coniogramme, Cryptogramma, and Llavea

Parkerioideae

• Acrostichum and Ceratopteris

Pteridoideae

• Actiniopteris, Anogramma, Austrogramme, Cerosora, Cosentinia, Gastoniella, Jamesonia (incl. Eriosorus and Nephopteris), Onychium, Pityrogramma, Pteris (incl. Neurocallis and Platyzoma), Pterozonium, Syngramma, Taenitis, and Tryonia

Vittarioideae

• Adiantum, Ananthacorus, Antrophyopsis, Antrophyum, Haplopteris, Hecistopteris, Polytaenium, Radiovittaria, Rheopteris, Scoliosorus, Vaginularia, and Vittaria

Stems

• Creeping or erect rhizomes

Leaves

• The leaves are almost always compound

Reproductive features

• Fronds have linear sori that are typically on the margins of the leaves and lack a true indusium, typically being protected by a false indusium formed from the reflexed margin of the leaf.

Saccolomataceae

• 1 genus (Saccoloma) with 18 species that are pantropical

Stems

• Terrestrial; rhizomes short-creeping to erect and trunk-like

Leaves

• Petioles each with an omega-shaped vascular strand; blades pinnate to decompound and lacking articulate hairs; veins free

Reproductive features

• Sori terminal on the veins; indusia pouch- or cup-shaped

Athyriaceae

• 3-6 genera (Anisocampium, Athyrium, Cornopteris, Deparia, Diplazium, and Pseudathyrium) with roughly 660 species

• This group has worldwide distribution, particularly the genus Athyrium

• Most species of Athyriaceae are medium-sized terrestrial or lithophytic ferns, growing in the understory of forests; less commonly aquatic

• The distribution and evolution of characters in the family are complex, and the genera have few constant features by which they can be identified.

Stems

• These ferns grow from various kinds of rhizome: short or long, creeping or erect, branched or not.

Leaves

• Pinnately-compound fronds

Reproductive features

• The sporangia have stalks two or three cells wide in the middle, and contain brown monolete spores

Blechnaceae

• 3 subfamilies with 24 genera (see below) with approximately 265 species

• Most members are ground-dwelling, some are climbers such as Stenochlaena

Woodwardioideae

• Anchistea, Lorinseria, and Woodwardia

Stenochlaenoideae

• Salpichlaena, Stenochlaena, and Telmatoblechnum

Blechnoideae

• Austroblechnum, Blechnidium, Blechnopsis, Blechnum, Brainea, Cleistoblechnum, Cranfillia, Diploblechnum, Doodia, Icarus, Lomaria, Lomaridium, Lomariocycas, Neoblechnum, Oceaniopteris, Parablechnum, Sadleria and Struthiopteris

Stems

• The chain ferns were formerly considered to belong to the Polypodiaceae.

Leaves

• The leaves are large and are once- or twice-divided, depending upon the species. The rachis is typically grooved on its upper side.

• ?many species is that the young opening fronds are usually tinged with red

Reproductive features

• Sori are elongate (i.e., non-circular) and arranged along the veins of the leaflets

• They are covered with an indusium and open toward the midvein or midrib.

• Sterile and fertile fronds can be similar or dimorphic

Cystopteridaceae

• 3 genera: Acystopteris with 3 species, Cystopteris with 14 species, and Gymnocarpium with 9 species

• Members are generally small or medium-sized ferns in forests and crevices

• This family is basal in the Aspleniineae (eupolypods II)

• Known as the Bladderferns, Brittleferns, and Oakferns

Stems

• These ferns generally have a slender, creeping rhizome under the surface of the ground may grow in rocky areas

• Rhizomes epigeous or more often subterranean, short- to more often long-creeping, occasionally suberect (Cystopteris), commonly branched, bearing scales and sometimes golden hairs similar to the root-hairs (e.g., C. protrusa)

• Rhizome scales lanceolate, clathrate or non-clathrate, the margins glandular or not, without distinct pubescence, entire to ciliate, the teeth when present not formed by two adjacent cells

Roots

• Roots blackish, wiry, inserted radially, non-proliferous;

Leaves

• Leaves green and not covered in mucilage during any stage of development, spirally arranged, monomorphic, bulbiferous in a few Cystopteris, closely spaced to distant, bearing scales and sometimes gland-tipped hairs, the scales sometimes reduced to filiform proscales (Cystopteris) or catenate hairs (Acystopteris); petioles stramineous throughout or proximally darkened, the base narrow, or conspicuously thickened and then starch-filled and persistent (trophopods), without conspicuous aerophores, without a proximal articulation, sometimes with golden hairs similar to the root hairs (e.g., C. moupinensis)

• Petiolar vascular bundles two, the bundles with hippocampiform-shaped xylem, distally uniting to form a single V-shaped bundle; laminae thin-herbaceous, 2–3-pinnate-pinnatifid (pinnate-pinnatifid in Cystoathyrium), broadest at the base or lanceolate, the apex non-conform, the leaf marginal cells differentiated into nodulose hyaline cells (Acystopteris, Cystopteris) or not (Gymnocarpium)

• Pinna axes distinctly articulate in Gymnocarpium, otherwise non-articulate, sulcate adaxially, lacking a free central ridge

• The rachis grooves continuous or not, the sulcus wall of the rachis continuing as a prominent ridge onto the sulcus wall of the costa or not

• Veins free, terminating at the leaf margin, the vein endings not differentiated

Reproductive features

• Sori dorsal along veins, not terminal, round or slightly elongate (Gymnocarpium), indusiate (Acystopteris, Cystopteris), or exindusiate (Gymnocarpium)

• Soral receptacle distinctly raised and hardened (Acystopteris, Cystopteris) or flat (Gymnocarpium)

• Indusia basal (Acystopteris, Cystopteris)

• Sporangia with stalks two or three cells wide in the middle

• Spores monolete, non-chlorophyllous, tan (Acystopteris) or brown, the perispore echinate, tuberculate, or with broad folds, the folds sometimes perforate

Diplaziopsidaceae

• 2 genera, Diplaziopsis with 3 species, and Homalosorus with 1 species: H. pycnocarpos

• Medium-to-large ferns, terrestrial or epipetric, which grow near streams in forested areas

• Species are found in east Asia, from China south to New Guinea and east into the Pacific

Stems

• The rhizomes are thick and decumbent to erect to suberect (Diplaziopsis, Diplazium flavoviride) or short-creeping (Homalosorus), commonly unbranched, bearing scales, and sometimes golden hairs similar to the root hairs (Homalosorus)

• Rhizome scales lanceolate, non-clathrate, the margins entire, non-glandular, without distinct pubescence; leaves green and not covered in mucilage during any stage of development, spirally arranged, monomorphic, non-bulbiferous, closely spaced, glabrous (Diplaziopsis) or with filiform proscales (Homalosorus)

Roots

• Roots fleshy, pale, inserted radially, non-proliferous

Leaves

• Petioles stramineous throughout or proximally darkened, thin, without a proximal thickening, conspicuous aerophores, or proximal articulation, sometimes with golden hairs similar to the root hairs (Homalosorus)

• Petiolar vascular bundles two, each with hippocampiform xylem, the bundles distally uniting to form a single V-shaped bundle; laminae soft-herbaceous, 1-pinnate, the apex conform (Diplaziopsis) or non-conform (Homalosorus), the leaf marginal cells differentiated into nodulose hyaline cells

• Pinna axes not articulate, sulcate adaxially, lacking a free central ridge

• The rachis grooves V-shaped, not continuous, the sulcus wall of the rachis continuing as a prominent ridge onto the sulcus wall of the costa; veins free (D. flavoviride, Homalosorus) or anastomosing toward the pinna margins (Diplaziopsis), the areoles without free included veinlets, usually terminating before the leaf margin, however some veins reaching the leaf margin in D. flavoviride and Homalosorus, the vein endings differentiated, slightly raised and expanded

Reproductive features

• Sori singular along one side of the vein, rarely paired back to back along the same vein, elongate, indusiate, not terminal

• Soral receptacle flat; indusia lateral, vaulted or essentially flat, glabrous or glandular (Diplaziopsis), opening along the lateral margin or sometimes rupturing irregularly (Diplaziopsis)

• Sporangia with stalks two or three cells wide in the middle; spores monolete, non-chlorophyllous, brown, the perispore folded with thin crests, the crests erose

Rhachidosoraceae

• 1 genus (Rhachidosorus) with 4-8 species

• Ferns from eastern and southeastern Asia, including Japan, the Philippines, and Sumatra

• Plants terrestrial

Stems

• Rhizomes creeping or short-creeping, not commonly branched, bearing scales

• Rhizome scales lanceolate, clathrate (with bars or latticed), the margins entire, without distinct pubescence

Roots

• Roots inserted radially, non-proliferous

Leaves

• Leaves green and not covered in mucilage during any stage of development, spirally arranged, monomorphic, not articulate to the rhizome, closely spaced, sparsely scaly

• Petioles reddish to stramineous throughout, narrow at the base, not forming trophopods, without conspicuous aerophores, without a petiolar articulation

• Petiolar vascular bundles two, each with hippocampiform xylem, the bundles distally uniting to form a single U-shaped bundle

• Laminae herbaceous, 2–3-pinnatepinnatifid, broadest at the base, the apex non-conform, the leaf marginal cells differentiated into nodulose hyaline cells

• Pinna axes not articulate, sulcate adaxially, lacking a free central ridge

• The rachis grooves U-shaped, continuous, the sulcus wall of the rachis continuing as a prominent ridge onto the sulcus wall of the costa, and then departing on the costule of the first basiscopic segment; veins free, terminating before the leaf margin, the vein endings not differentiated

Reproductive features

• ?sori dorsal along

• veins, not terminal, elongate, indusiate; soral receptacle flat; indusia

• lateral, non-glandular; sporangia with stalks two or three

• cells wide in the middle; spores monolete, non-chlorophyllous,

• brown, the perispore echinate, tuberculate, or with broad folds,

• the folds sometimes perforate;

Didymochlaenaceae

• 1 species, Didymochlaena truncatula, also known as the mahogany maidenhair

• This family is basal in the Polypodiineae, but this genus is sometimes placed within the Hypodematiaceae

Stems

• ?

Leaves

• ?

Reproductive features

• Elongated sori are present and different from those of any other eupolypods I, but similar to those of some Aspleniaceae or Athyriaceae of eupolypods II

• This species has dimidiate pinnules

Dryopteridaceae

• This family of wood ferns contains 3 subfamilies, 24 genera, and ~1,700 species

• 70% of species are in 4 of these genera: Elaphoglossum (600+), Polystichum (260), Dryopteris (225), and Ctenitis (150)

Polybotryoideae

• Cyclodium, Maxonia, Olfersia, Polybotrya, Polystichopsis, Stigmatopteris, and Trichoneuron

Elaphoglossoideae

• Arthrobotrya, Bolbitis, Elaphoglossum, Lastreopsis, Lomagramma, Megalastrum, Mickelia, Parapolystichum, Pleocnemia, Rumohra, and Teratophyllum,

Dryopteridoideae

• Arachniodes, Ctenitis, Cyrtomium, Dryopteris, Phanerophlebia, and Polystichum

Stems

• Rhizomes are often stout, creeping, ascending, or erect, and sometimes scandent or climbing, with nonclathrate scales at apices

Leaves

• Fronds are usually monomorphic, less often dimorphic, or sometimes scaly or glandular, but less commonly hairy

• Petioles have numerous round, vascular bundles arranged in a ring, or rarely as few as three; the adaxial bundles are the largest

• Veins are pinnate or forking, free to variously anastomosing; the areoles occur with or without included veinlets

Reproductive features

• Sori are usually round, acrostichoid (covering the entire abaxial surface of the lamina) in a few lineages

• Usually indusiate, or sometimes exindusiate

• Indusia, when present, are round-reniform or peltate

• Sporangia have three-rowed, short to long stalks

• Spores are reniform, monolete, perine or winged

Hypodematiaceae

• 2 genera: Hypodematium with ~20 species, and Leucostegia with 2 species (L. pallida and L. truncata)

Stems

• ?

Leaves

• ?

Reproductive features

• ?

Lomariopsidaceae

• Four genera: Cyclopeltis, Dracoglossum, Dryopolystichum, Lomariopsis); ~70 species.

Stems

• Rhizomes creeping or sometimes climbing (plants hemi-epiphytic)

Leaves

• Petioles with round vascular bundles arranged in a gutter-shape;

• Blades 1-pinnate, pinnae entire or crenate, often articulate, auriculate in some genera; veins free, ± parallel or pinnate

Reproductive features

• Sori discrete, round, and with round-reniform to reniform indusia, or exindusiate, or sporangia acrostichoid and the fronds dimorphic

• Spores bilateral, monolete, variously winged or ornamented

Nephrolepidaceae

• 1 genus (Nephrolepis) with ~20 species

• The genus is commonly referred to as macho ferns or swordferns, but N. exaltata is known as Boston fern

• Most species can grow as terrestrials or epiphytes

Stems

• Complex rhizome system with both erect, frond-bearing stems and creeping, root-bearing and sometimes tuber-forming runners

• The rhizomatous runners can grow subterraneously, on the surface or in aerial thickets

Leaves

• Simply pinnate fronds with articulate pinnae

Reproductive features

• The sori are terminal on the acroscopic vein-branch, in a medial to marginal position, and indusiate with a reniform to lunulate or linear indusium

Oleandraceae

• 1 genus (Oleandra) with 15 species

Stems

• Most are erect ground ferns or scandent epiphytes that start from the ground.

Leaves

• The lamina (leafy area of the fronds) are simple or pinnate, and the individual pinnae are articulate to the rachis.

Reproductive features

• The sporangia are contained in discrete round sori in a single row on either side of the midrib of the fronds

Onocleaceae

• 4 genera with 5 species: Matteuccia struthiopteris, Onoclea sensibilis, Onocleopsis hintonii, Pentarhizidium orientale & P. intermedium)

Stems

• The rhizomes are long- to short-creeping to ascending, and sometimes stoloniferous (Matteuccia and Onocleopsis)

• The leaves are strongly dimorphic and the petioles have two vascular bundles uniting distally into a gutter-shape

• The blades are pinnatifid or pinnate-pinnatifid

• The veins are free or anastomosing, lacking included veinlets

Leaves

• Strongly dimorphic fronds, with the fertile fronds different from the sterile fronds.

Reproductive features

• The spores are reniform, brownish to green

• The sori are enclosed (sometimes tightly) by reflexed laminar margins, also with membranous, often fugacious true indusia

Polypodiaceae

• 5 subfamilies; 65 genera; ~1,650 species

Loxogrammoideae

• Dictymia and Loxogramme

Drynarioideae

• Aglaomorpha, Arthromeris, Gymnogrammitis, Paraselliguea, Polypodiopteris, and Selliguea

Platycerioideae

• Platycerium and Pyrrosia

Microsoroideae

• Goniophlebium, Lecanopteris, Lemmaphyllum, Lepidomicrosorium, Lepisorus, Leptochilus, Microsorum, Neocheiropteris, Neolepisorus, Paragramma, Thylacopteris and Tricholepidium

Polypodioideae

• Campyloneurum, Microgramma, Niphidium, Pecluma, Phlebodium, Pleopeltis, Pleurosoriopsis, Polypodium and Serpocaulon

Grammitidoideae

• Acrosorus, Adenophorus, Alansmia, Archigrammitis, Ascogrammitis, Calymmodon, Ceradenia, Chrysogrammitis, Cochlidium, Ctenopterella, Dasygrammitis, Enterosora, Galactodenia, Grammitis, Lellingeria, Leucotrichum, Lomaphlebia, Luisma, Melpomene, Micropolypodium, Moranopteris, Mycopteris, Notogrammitis, Oreogrammitis, Prosaptia, Radiogrammitis, Scleroglossum, Stenogrammitis, Terpsichore, Themelium, Tomophyllum, Xiphopterella, and Zygophlebia

Stems

• ?

Leaves

• ?

Reproductive features

• ?

Thelypteridaceae

• 2 subfamilies; 30 genera; ~900 species, called the marsh ferns

• The ferns are terrestrial, with the exception of a few which are lithophytes (grow on rocks)

• The bulk of the species are tropical, although there are a number of temperate species

• Cretaceous-present

Phegopteridoideae

• Macrothelypteris, Phegopteris, and Pseudophegopteris

Thelypteridoideae

• Amauropelta, Amblovenatum, Ampelopteris, Chingia, Christella, Coryphopteris, Cyclogramma, Cyclosorus, Glaphyropteridopsis, Goniopteris, Meniscium, Menisorus, Mesophlebion, Mesopteris, Metathelypteris, Nannothelypteris, Oreopteris, Parathelypteris, Plesioneuron, Pneumatopteris, Pronephrium, Pseudocyclosorus, Sphaerostephanos, Stegnogramma, Steiropteris, Thelypteris, and Trigonospora

Stems

• These ferns typically have creeping rhizomes.

Leaves

• The fronds are simply pinnate to pinnate-pinnatifid

• There is either no frond dimorphism or only mild dimorphism, either open venation or very simple anastomosing

Reproductive features

• The sori are mostly reniform in shape and have indusia, except for the Phegopteridoideae