Mangrove fern Acrostichum aureum, photographed by Andrés Hernández.

Belongs within: Polypodiales.
Contains: Cheilanthes, Vittariaceae, Adiantum, Pteris.

The Pteridaceae is a cosmopolitan family of ferns with abaxial sori lacking indusia or protected by a reflexed or revolute leaf margin (M. D. Windham). The molecular phylogenetic analysis of ferns by Schneider et al. (2004) suggested that the Pteridaceae may be paraphyletic with regard to the Vittariaceae. The Pteridaceae also includes the aquatic genus Ceratopteris, as well as the semi-aquatic Acrostichum. Cheilanthes, the lip ferns, is a cosmopolitan genus of ferns found growing on rocks in warm, dry regions.

Characters (from M. D. Windham): Plants perennial (occasionally annual), on rock or terrestrial, of small (rarely large) stature . Stems compact to creeping, branched or unbranched, dictyostelic, bearing hairs and/or scales. Leaves monomorphic to dimorphic, circinate or noncircinate in bud . Petioles usually with persistent scales proximally, lacking spines; vascular bundles 1-several, roundish or crescent-shaped in cross section. Blades 1-6-pinnate, without laminar buds . Indument on petioles, rachises, costae, and blades, rarely absent or commonly of hairs, glands, and/or scales, occasionally of white or yellow farina. Veins pinnate or parallel in ultimate segments of blades, simple or forked, free or infrequently anastomosing in complex patterns. Sori borne abaxially on veins, often confluent with age and forming a continuous submarginal band, or sporangia densely covering abaxial surface (acrostichoid); receptacle not or only slightly elevated. Indusia (when present) formed by reflexed, recurved, or revolute leaf margin (false indusium). Sporangia stalk of 2–3 rows of cells; annulus vertical, interrupted by stalk; spores 64 or 32 (rarely 16) per sporangium. Spores all 1 kind, brown, black, or gray (rarely yellow), globose to globose-tetrahedral or trigonal, occasionally with prominent equatorial ridge, trilete, or trigonal, variously ornamented (usually cristate or rugose). Gametophytes green, aboveground, obcordate to reniform, sometimes asymmetric, usually glabrous (glandular-farinose in Notholaena); archegonia and antheridia borne on abaxial surface, antheridia 3-celled.

<==Pteridaceae [Adiantaceae, Sinopteridaceae]
    |  |    |--B. ehrenbergianaSS04
    |  |    `--B. pedataL54
    |  `--+--VittariaceaeSS04
    |     `--+--AdiantumSS04
    |        `--Hewardia regiaSS04
          |  `--CeratopterisSS04
          |       |--C. richardiiSL05
          |       `--C. thalictroides (Linnaeus) Brongniart 1822I88 (see below for synonymy)
               |--A. aureumSS04 [incl. A. fraxinifoliumB78, A. inaequaleBS-V28, A. speciosumBS-V28]
               |--A. conforme [=Elaphoglossum conforme]B78
               |--A. danaeifoliumSS04
               |--A. drynarioidesT-W89
               |--A. latifoliumV97
               |--A. minus [=Gymnopteris minor]T-W89
               |--A. nervosum Bory de Saint-Vincent 1828BS-V28
               |--‘Osmunda’ peltataV97
               |--A. pteroides [=Neurosoria pteroides]B78
               |--A. repandumB78
               |--A. rigidum [=Photinopteris rigida]T-W89
               |--A. scandensB78
               |--A. sorbifolium [incl. A. brightiae, Lomariopsis brightiae, L. leptocarpa]B78
               |--A. spicatumB78 [=Hymenolepis spicataB78, Lomaria spicataBS-V28]
               |--A. subrepandumT-W89
               |--A. trifoliatumBS-V28
               |--A. variabileT-W89
               `--A. villosumV97
Pteridaceae incertae sedis:
    |--A. jonesii [=Cheilanthes jonesii]H93
    `--A. limitanea [=Cheilanthes limitanea, Notholaena limitanea]H93
  Anopteris hexagoniaJ87
  Gymnopteris Bernhardi 1799YS03
    `--G. vestita (Bedd.) Underwood 1902 (see below for synonymy)I88
  Notholaena Brown 1810YS03 [=Chrysochosma (Kümm.) Pichi Sermolli 1989YS03, NothochlaenaB78]
    |--*N. trichomanoides (Linnaeus) Brown 1810 [=Pteris trichomanoides]YS03
    |--N. californica [incl. N. californica ssp. nigrescens]H93
    |--N. fragilis [incl. Cheilanthes fragillima]B78
    `--N. parryiGCR98
  Paragymnopteris Shing 1993YS03
    `--*P. marantae (Linnaeus) Shing 1993 (see below for synonymy)YS03
  Pellaea Link 1841A61
    |--P. andromedifolia [incl. P. andromedifolia var. pubescens, P. andromedifolia var. rubens]H93
    |--P. atropurpureaSL05
    |--P. brachypteraH93
    |--P. breweriH93
    |--P. bridgesiiH93
    |--P. falcata (Br.) Fée 1850-1852 (see below for synonymy)A61
    |    |--P. f. var. falcataH90
    |    `--P. f. var. nanaH90
    |--P. × glaciogena [P. bridgesii × P. mucronata]H93
    |--P. mucronataH93
    |    |--P. m. var. mucronataH93
    |    `--P. m. var. californica [incl. P. compacta]H93
    |--P. paradoxaH90 [=Adiantum paradoxumB78, Pteris paradoxaB78; incl. Platyloma browniiB78]
    |--P. rotundifolia (Forster) Hook. 1858 (see below for synonymy)A61
    `--P. truncataH93
  Paesia St. Hilaire 1833A61
    `--P. scaberula (Rich.) Kuhn 1882 (see below for synonymy)A61
  Paraceterach muelleriLK14
    |--A. californicaH93
    |--A. carlotta-halliae [=Cheilantes carlotta-halliae]H93
    `--A. densa [=Onychium densum]H93
  Astrolepis cochisensis [=Cheilanthes sinuata var. cochisensis]H93
    |--C. acrostichoidesH93
    |--C. brunoniana Wallich ex Hooker & Greville 1829 [incl. C. crispa (Linnaeus) Brown in Richards 1823]I88
    |--C. cascadensisH93
    `--C. stelleri (Gmelin) Prantl 1882 (see below for synonymy)I88
    |--P. pallida [=Pityrogramma triangularis var. pallida]H93
    `--P. triangularis [=Pityrogramma triangularis]H93
         |--P. t. ssp. triangularis [incl. P. triangularis ssp. semipallida]H93
         |--P. t. ssp. maxonii [=Pityrogramma triangularis var. maxonii]H93
         `--P. t. ssp. viscosa [=Pityrogramma triangularis var. viscosa]H93
Nomina nuda: Pellaea saportana Squinabol 1889S89

Ceratopteris thalictroides (Linnaeus) Brongniart 1822I88 [=Acrostichum thalictroides Linnaeus 1753I88; incl. Parkeria pteridioidesB78, A. siliquosum Linnaeus 1753I88, Ceratopteris siliquasa (Linnaeus) Copel. 1935I88]

Cryptogramma stelleri (Gmelin) Prantl 1882 [=Pteris stelleri Gmelin 1768, Pellaea stelleri (Gmelin) Bedd. 1883; incl. Pe. gracilis Hooker 1858]I88

Gymnopteris vestita (Bedd.) Underwood 1902 [=Syngramma vestita Bedd. 1883, Grammitis vestita Wallich ex Bedd. 1883 (n. n.), Gymnogramma vestita]I88

Paesia scaberula (Rich.) Kuhn 1882 [=Pteris scaberula Rich. 1832, Allosurus scaberulus Presl 1836; incl. Pt. microphylla Cunn. 1837]A61

*Paragymnopteris marantae (Linnaeus) Shing 1993 [=Acrostichum marantae Linnaeus 1753, Notholaena marantae (Linnaeus) Brown 1810, Paraceterach marantae (Linnaeus) Tryon 1986]YS03

Pellaea falcata (Br.) Fée 1850-1852 [=Pteris falcata Br. 1810, Platyloma falcata Smith 1841; incl. Pt. seticaulis Hook. 1840]A61

Pellaea rotundifolia (Forster) Hook. 1858 [=Pteris rotundifolia Forster 1786, Allosurus rotundifolius Kunze 1856, Platyloma rotundifolia Smith 1841]A61

*Type species of generic name indicated


[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[B78] Bentham, G. 1878. Flora Australiensis: A description of the plants of the Australian Territory vol. 7. Roxburghiaceae to Filices. L. Reeve & Co.: London.

[BS-V28] Bory de Saint-Vincent, J. B. 1828. Voyage Autour du Monde, Exécuté par Ordre du Roi, Sur la Corvette de Sa Majesté, La Coquille, pendant les années 1822, 1823, 1824 et 1825. Botanique. Cryptogamie. Arthus Bertrand: Paris.

[GCR98] Gratani, L., M. F. Crescente & G. Rossi. 1998. Photosynthetic performance and water use efficiency of the fern Cheilanthes persica. Photosynthetica 35: 507–516.

[H90] Harden, G. J. (ed.) 1990. Flora of New South Wales vol. 1. New South Wales University Press.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[I88] Iwatsuki, K. 1988. An enumeration of the pteridophytes of Nepal. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 231–339.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.

[L54] Lötschert, W. 1954. Beitrag zur Pteridophyten-Flora von Mittel-Amerika. Senckenbergiana Biologica 35 (1–2): 109–119.

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[SS04] Schneider, H., E. Schuettpelz, K. M. Pryer, R. Cranfill, S. Magallón & R. Lupia. 2004. Ferns diversified in the shadow of angiosperms. Nature 428: 553–557.

[SL05] Small, R. L., E. B. Lickey, J. Shaw & W. D. Hauk. 2005. Amplification of noncoding chloroplast DNA for phylogenetic studies in lycophytes and monilophytes with a comparative example of relative phylogenetic utility from Ophioglossaceae. Molecular Phylogenetics and Evolution 36: 509–522.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 73–76.

[T-W89] Tenison-Woods, J. E. 1889. On the vegetation of Malaysia. Proceedings of the Linnean Society of New South Wales, series 2, 4 (1): 9–106, pls 1–9.

[V97] Ventenat, C. 1797. Extrait d’un mémoire sur les fougeres, de M. Smith. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (9): 71–72.

[YS03] Yatskievych, G., & A. R. Smith. 2003. Typification of Notholaena R. Br. (Pteridaceae). Taxon 52: 331–336.

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