Mangrove fern Acrostichum aureum, photographed by Andrés Hernández.

Belongs within: Pteridales.
Contains: 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]
    |--+--Bommeria SS04
    |  |    |--B. ehrenbergiana SS04
    |  |    `--B. pedata L54
    |  `--+--Vittariaceae SS04
    |     `--+--Adiantum SS04
    |        `--Hewardia regia SS04
    `--+--Pteris SS04
       `--+--Acrostichum SS04
          |    |--A. aureum SS04
          |    |--A. danaeifolium SS04
          |    `--A. speciosum WF01
          `--+--Magnastriatites SS04
             `--Ceratopteris SS04
                  |--C. richardii SL05
                  `--C. thalictroides (Linnaeus) Brongniart 1822 (see below for synonymy) I88

Pteridaceae incertae sedis:
  Argyrochosma YS03
  Anopteris hexagonia J87
  Gymnopteris Bernhardi 1799 YS03
    `--G. vestita (Bedd.) Underwood 1902 (see below for synonymy) I88
  Notholaena Brown 1810 [=Chrysochosma (Kümm.) Pichi Sermolli 1989] YS03
    |--*N. trichomanoides (Linnaeus) Brown 1810 [=Pteris trichomanoides] YS03
    `--N. parryi GCR98
  Paragymnopteris Shing 1993 YS03
    `--*P. marantae (Linnaeus) Shing 1993 (see below for synonymy) YS03
  Pellaea Link 1841 A61
    |--P. atropurpurea SL05
    |--P. falcata (Br.) Fée 1850-1852 (see below for synonymy) A61
    `--P. rotundifolia (Forster) Hook. 1858 (see below for synonymy) A61
  Cheilanthes Swartz 1806 A61 [incl. Aleuritopteris I88]
    |--C. albomarginata Clarke 1880 [=C. farinosa var. albomarginata (Clarke) Bedd. 1892] I88
    |--C. anceps Blanford 1886 I88
    |--C. angustifolia L54
    |--C. aurea L54
    |--C. austrotenuifolia G04
    |--C. brownii LK14
    |--C. caudata LK14
    |--C. chrysophylla Hooker 1852 [=C. farinosa var. chrysophylla (Hooker) Clarke 1880] I88
    |--C. contigua LK14
    |--C. dalhousiae Hooker 1852 (see below for synonymy) I88
    |--C. deltoidea KLN06
    |--C. distans (Brown) Mett. 1859 [=Notholaena distans Brown 1810] A61
    |--C. duthiei Baker 1891 I88
    |--C. farinosa (Forskål) Kaulf. 1824 (see below for synonymy) I88
    |--C. fragillima LK14
    |--C. fragrans D30
    |--C. kaulfussii L54
    |--C. lanosa SL05
    |--C. lasiophylla G04
    |--C. notholaenoides J87
    |--C. nudiuscula LK14
    |--C. odora C74
    |--C. persica GCR98
    |--C. pteridioides PT98
    |--C. pumilio LK14 [=Notholaena pumilio Brown 1810 YS03]
    |--C. pyramidalis L54
    |--C. rufa Don 1825 I88
    |--C. setigera Blume 1828 A61 (see below for synonymy)
    |--C. sieberi Kunze in Lehm. 1846 [incl. C. erecta Colenso 1896] A61
    |--C. subvillosa Hooker 1852 [=Aleuritopteris subvillosa (Hooker) Ching 1941] I88
    |--C. tamburii (Hooker) Moore 1861 [=Pellaea tamburii Hooker 1858] I88
    |--C. tenuifolia (Burmann) Swartz 1806 I88 (see below for synonymy)
    `--C. vellea PT98 [=Notholaena vellea Brown 1810 YS03]
  Neurocallis SS04
  Paesia St. Hilaire 1833 A61
    `--P. scaberula (Rich.) Kuhn 1882 (see below for synonymy) A61
  Paraceterach muelleri LK14
  Platyzoma microphyllum LK14

Nomina nuda: Pellaea saportana Squinabol 1889 S89

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

Cheilanthes dalhousiae Hooker 1852 [=C. farinosa var. dalhousiae (Hooker) Clarke 1880, Leptolepidium dalhousiae] I88

Cheilanthes farinosa (Forskål) Kaulf. 1824 [=Pteris farinosa Forskål 1775; incl. C. dealbata Don 1825 non Pursh 1814, C. grisea Blanford 1886, Aleuritopteris grisea (Blanford) Panigr. 1961] I88

Cheilanthes setigera Blume 1828 A61 [incl. Dryopteris setigera A61, Thelypteris setigera (Blume) Ching 1936 C49]

Cheilanthes tenuifolia (Burmann) Swartz 1806 I88 [=Trichomanes tenuifolium Burmann 1768 I88; incl. C. kirkii Armstrong 1881 (n. n.) A61, C. venosa Colenso 1893 A61]

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

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

[C74] Coineau, Y. 1974. Éléments pour une monographie morphologique, écologique et biologique des Caeculidae (Acariens). Mémoires du Muséum National d'Histoire Naturelle, nouvelle série, Série A, Zoologie 81: 1–299, 224 pls.

[C49] Crookes, M. W. 1949. A revised and annotated list of New Zealand Filicinae. Transactions and Proceedings of the Royal Society of New Zealand 77 (2): 209–225.

[D30] Druce, G. C. 1930. Account of a botanical tour in Cyprus. Proceedings of the Linnean Society of London 141: 50–52.

[G04] Gibson, N. 2004. Flora and vegetation of the Eastern Goldfields Ranges: part 6. Mt Manning Range. Journal of the Royal Society of Western Australia 87 (2): 35–47.

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

[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.

[KLN06] Klopper, R. R., P. Lemmer & J. Nel. 2006. Pteridophyta: Pteridaceae. Cheilanthes deltoidea, a new locality in Gauteng, South Africa. Bothalia 36 (2): 173–174.

[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.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[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.

[WF01] Woinarski, J. C. Z., A. Fisher, K. Brennan, I. Morris & R. Chatto. 2001. Patterns of bird species richness and composition on islands off Arnhem Land, Northern Territory, Australia. Austral Ecology 26: 1–13.

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

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