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Incl. Sphaerostephanaceae

Terrestrial or rupestral ferns, sometimes growing along swamps or marshes; rhizome erect to long-creeping, dictyostelic, bearing scales. Fronds mostly long-stipitate, the stipe with 2 vascular strands at the base uniting upwards to form one of the U-shape. Fronds mostly pinnate with crenate to pinnatifid pinnae, uncommonly simple or bipinnate, glabrous or with various hairs or glands, venation in each pinna of pinnate groups about a costule in each lobe, free, terminating at the margin or just short of it, often the basal veins of each group uniting to form a single excurrent vein that terminate in the sinus membrane. Sori borne on the adaxial surface of the veins, round or sometimes slightly elongate, with a +/- reniform indusium or not; sporangia often bearing hairs or glands; annulus longitudinal, interrupted; spores nearly always monolete.


A predominently tropical family of c. 30 genera (or much less depending on the author) encompassing more than 1000 species. In Papuasia there are 17 genera in the narrow sense, with more than 220 species.


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

Holttum, R.E. 1969. Studies in the Thelypteridaceae. The genera Phegopteris, Pseudophegopteri and Macrothelypteris. Blumea 17: 5 - 32.

Holttum, R.E. 1971. Studies in the family Thelypteridaceae III. A new system of genera in the old world. Blumea 19: 17 - 52, f. 1 - 19.

Holttum, R.E.- 1972. Studies in the family Thelypteridaceae IV. The genus Pronephrium Presl. Blumea 20: 105 - 126.

Holttum, R.E. 1973. Studies in the Thelypteridaceae V. The genus Pneumatopteris Nakai. Blumea 21: 293 - 325.

Holttum, R.E. 1974. Studies in the family Thelypteridaceae VII. The genus Chingia. Kalikasan 3: 13 - 28.

Holttum, R.E. 1975. Studies in the Thelypteridaceae VIII. The genera Mesophlebion and Plesioneuron. Blumea 22: 223 - 250.

Holttum, R.E. 1976. Studies in the Thelypteridaceae X. The genus Coryphopteris. Blumea 23: 18 - 47.

Holttum, R.E. 1976. The genus Christella Leveille sect. Christella. Studies in the Thelypteridaceae, XI. Kew Bull. 31: 293 - 339.

Holttum, R.E. 1977. Studies in the Thelypteridaceae XII. The genus Amphineuron Holttum. Blumea 22: 205 - 218.

Holttum, R.E. 1977. The family Thelypteridaceae in the Pacific and Australasia. Allertonia 1: 169 - 243, f. 1 - 10.

Holttum, R.E. 1982. Thelypteridaceae. Fl. Males. ser. 2 1: 331 - 560, f. 1 - 20.

Holttum, R.E., Sen, U. & Mittra, D. 1970. Studies in the family Thelypteridaceae II. A comparitive study of the type-species of Thelypteris Schmidel, Cyclosorus Link, and Ampelopteris Kunze. Blumea 18: 195 - 215, f. 1 - 112.



Costae not grooved on the upper surface; tips of veins not reaching the margin


Costae grooved on the upper surface; free veins all reaching the margin



Fronds bipinnate

Macrothelypteris (1)

Fronds simply pinnate, sometimes pinnae very deeply lobed



Sori indusiate

Metathelypteris (1)

Sori exindusiate

Pseudophegopteris (3)


Fronds simple, or pinnate with the lower pinnae not decresent (small basal pinnae are inconstantly present in Amphineuron)


Pinnate with the lower pinnae gradually reduced, or with an abrupt change to small pinnae at the base of the frond



Rhizome long-creeping in wet or swampy ground; broad flat scales present on the lower surface of costae


Not this combination of characters



Veins free

Thelypteris (1)

Veins anastomosing



Fronds not proliferous; sori indusiate

Cyclosorus (1)

Fronds with proliferating buds along the rachis; sori exindusiate

Ampelopteris (1)


Caudex erect; sporangia unstalked, lacking hairs and glands; some sessile resiniferous glands present on the lower surface of the lamina in most species; plants of mountain ridges

Coryphopteris (c. 22)

Not this combination of characters



Caudex massive, erect; scales narrow, rigid, brittle; sori close to costules, exindusiate or with a very small indusium

Chingia (5)

Not this combination of characters



Veins free


Veins anastomosing



Basal basiscopic vein of each group arising from the costa below the attachment of its costule; a red gland present at the ends of the hairs on the sporangium-stalk

Mesophlebion (3)

Basal basiscopic vein of each group not always thus arising; hairs on sporangium-stalk otherwise



Rhizome long-creeping; no glands or hairs on sporangia


Rhizome short-creeping or erect; sporangia in some cases with glands or hairs



Rhizome slender; pinnae rarely over 6cm long

Parathelypteris (1)

Rhizome 5-7mm diameter; pinnae 10cm or more long

Christella (10)


Bases of lower pinnae not greatly narrowed


Bases of lower pinnae much narrowed

Amphineuron (7)


Pinnae rigid, commonly at least 10cm long and 1cm wide

Plesioneuron (c. 37)

Pinnae thin, to 3cm long, rarely 1cm wide



Sporangia bearing glands

Pronephrium (c. 13)

Sporangia lacking glands

Christella (10)


Spores coarsely tuberculate or ridged; sporangia lacking glands


Spores with a thin wing and cross-wings, or many small thin wings



Basal pinnae much narrowed at their bases

Amphineuron (7)

Basal pinnae not much narrowed at their bases, which are in most cases auricled on the acroscopic side

Christella (10)


Pinnae or simple fronds subentire; lower surface between veins often pustular

Pronephrium (c. 13)

Pinnae deeply lobed; lower surface between veins not pustular

Sphaerostephanos (c. 62)


Rhizome slender, long-creeping; many lower pinnae gradually reduced; some septate acicular hairs on lower surface

Parathelypteris (1)

Not this combination of characters



Caudex erect; pinnae to c. 5cm long, the lower ones gradually reduced; sporangia sessile, lacking hairs or glands

Coryphopteris (c. 22)

Not this combination of characters



Basal scales on stipe in almost all cases broad and thin; lamina between veins on upper surface almost always smooth and glabrous, lower surface of costae never densely hairy; glandular hairs on lower surface, if present, minute and colourless

Pneumatopteris (c. 24)

Basal scales narrow; upper surface between the veins often bearing hairs and or glands, lower surface of costae usually with copious hairs; sessile, spherical or elongate glands often present



Body of sporangium lacking hairs or glands; an elongate unicellular glandular hair on the sporangium-stalk, similar hairs sometimes also on the lower surface of the veins; spores coarsely tuberculate or ridged

Christella (10)

Body of sporangium usually with sessile spherical glands or setae; hairs on sporangium stalks consisting of several cells; spores with many small thin wings or with a continuous wing with cross-wings; sessile spherical glands present on the lower surface of pinnae in many species.

Sphaerostephanos (c. 62)


This is the most finely divided treatment of the family, following the concepts of Holttum. At the opposite extreme other authors include all species in a single large genus, Thelypteris, and accommodate the variation in a large number of infrageneric taxa.

Updated November 1999 by Jim Croft (