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Moderate to large terrestrial ferns, rhizome mostly erect, less often creeping, mostly radial, dictyostelic, the apex bearing non-clathrate non-peltate scales, often with marginal teeth. Fronds long-stipitate, the stipes not articulate to the rhizome, with 3 or more vascular strands arranged in a simple ring, or in a more complex arrangement, less often with 2 strands not uniting upwards, lamina pinnate to pinnately decompound, less often simple, axes generally not grooved or if grooved the grooves not open to admit those of a higher or lower order, pinnate sometimes jointed to the rachis, veins free, mostly forked, or anastomosing with or without free include veinlets; sometimes marked dimorphism between sterile and fertile fronds. Sori mostly remote from the margin, dorsal or terminal on the veins, small, round or oblong, mostly indusiate, the indusium peltate or reniform, sometimes sori acrostichoid, the sporangia completely covering the lower surface of contracted fertile leaflets; annulus longitudinal, interrupted; spores bilateral, monolete, usually with a variously ornamented perispore.


A widespread family of up to 27 genera in the warmer parts of the globe. In Papuasia there are 12 genera with c. 60 species.


Copeland, E.B. 1949. Aspidiaceae of New Guinea. Philip. J. Sci. 78: 389 - 475.

Holttum, R.E. 1951. The fern genus Pleocnemia Presl. Reinwardtia 1: 171 - 189.

Holttum, R.E. 1951. The fern genus Arcypteris Underwood (Dictyopteris Presl sensu Fee. Reinwardtia 1: 191 - 196.

Holttum, R.E. 1974. The fern genus Pleocnemia. Kew Bull. 29: 341 - 357.

Holttum, R.E. 1975. The genus Heterogonium Presl. Kalikasan 4: 205 - 231.

Holttum, R.E. 1981. The fern genus Tectaria Cav. in Malaya. Gard. Bull. Sing. 34: 132 - 147.

Holttum, R.E. 1984. Studies in the fern genera allied to Tectaria. III. Aenigmopteris and Ataxipteris, two new genera allied to Tectaria Cav., with comments on Psomiocarpa Presl. Blumea 30: 1 - 11.

Holttum, R.E. 1985. Studies in the fern genera allied to Tectaria Cav. IV. The genus Ctenitis in Asia, Malesia and the Western Pacific. Blumea 31: 1- 38.

Holttum, R.E. & Edwards, P.J. (in press. Dryopsis, a new genus of ferns. Kew Bull.

Tindale, M.D. 1965. A monograph of the genus Lastreopsis Ching. Contrib. N.S.W. Nat. Herb. 3: 249 - 339.



A tooth present in each sinus between the lobes of the lamina, raised out of the plane of the lamina


No tooth present in the sinuses between the lobes of the lamina



Vascular bundles in the stipe numerous, not in a simple ring; veins anastomosing


Vascular bundles in the stipe in a simple ring; veins free

Pteridrys (2)


Veins anastomosing in a single series of areoles along the costae and costules

Pleocnemia (c. 7)

Veins more copiously anastomosing

Arcypteris (3)


Fertile leaflets acrostichoid or with linear, marginal sori, very much contracted as compared with the sterile leaflets


Fertile leaflets not acrostichoid, not or only slightly contracted as compared with sterile, with discrete, roundish sori



Fronds broadly deltoid, the basal basiscopic lobe of the basal pinnae the largest; a dormant bud present at the apex of the stipe

Stenosemia (1)

Fronds elliptic or oblong, the basal basiscopic lobe of the basal pinnae not enlarged; no dormant buds



Sterile fronds simple to deeply pinnatifid, or the upper half of the frond with broadly adnate, decurrent lobes, the margins +/- entire; the sori with a scariose, reflexed inner indusium, sometimes very indistinct

Hemmigramma (2)

Sterile fronds with at least 1 pair of free pinnae, usually several pairs, the pinnae crenate or lobed, exindusiate

Heterogonium (4)


Two vascular bundles in stipe; scales confined to the swollen base of the stipe (Athyriaceae)

Hypodematium (1)

Several vascular bundles in stipe; scales +/- scattered on lower portion of stipe



Veins free


Veins anastomosing



Basal basiscopic vein of each group arising from its own costule


Basal basiscopic vein of each group arising directly from the costa



Indusium peltate

Dryopolystichum (1)

Indusium reniform or absent



Fronds usually longer than wide; basiscopic margins of lamina lobes not thickened


Fronds usually about as long as wide; basiscopic margins of lamina lobes not thickened

Lastreopsis (1-2)


Thick hairs present between the veins on upper surface of lamina

Aenigmopteris (1)

Thick hairs lacking (thin-walled cells with prominent septa may be present)

Ctenitis (5)


Basal pinnae with the basal basiscopic lobes the longest

Tectaria (c.27)

Basal pinnae with basal basiscopic lobes or pinnules shorter than the middle lobes or pinnules

Heterogonium (4)


Many free veins in areoles

Tectaria (c. 27)

Free veins in areoles few or absent



Basal pinnae deeply lobed, the basal basiscopic lobe not the largest; few free veins in areoles

Heterogonium (4)

Basal pinnae pinnate at least towards the base, the basal basiscopic the longest

Arcypteris (3)


Some of the genera in this family are not readily separated. (Holttum 1974) chose to combine Arcypteris under Pleocnemia. Tectaria and Ctenitis are fairly heterogeneous and may include other genera or parts of other genera. This problem is being resolved by Holttum who has recently described Aenigmopteris and proposed a new genus Dryopsis (Holttum 1984 and in press).

The genus Hemigramma has been united with Tectaria (Price 1974) based on Philippine species which lack indusia; analagous situations of combining other tectarioid genera with acrostichoid or linear marginal sori with Tectaria exist in Ceylon and South America. However, the Papuasian species are indusiate; if retained separate from Hemigramma and Tectaria they will require a new generic name. Holttum (pers. comm.) is proposing the new genus Chlamydogramme for the Papuasian species that have been included in Hemmigramma.

Hypodematium has been included here by some authors, but on balance of characters it seems better placed in Athyriaceae.

Dryopteridaceae is often united with Hypoderriaceae and sometimes Lomariopsidaceae, Thelypteridaceae and Athyriaceae, in a large and complex family, the Aspidiaceae.

Updated November 1999 by Jim Croft (