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A Revised and Annotated List of New Zealand Filicinae

[Read before the Auckland Institute, March, 1947; received by Editor, April 12, 1947; issued separately February, 1949.]

A revision of New Zealand fern names is long overdue. It is hoped that the following list will give some indication of the present position. It was originally prepared prior to the publication of Professor E. B. Copeland's Genera Filicum (Chronica Botanica, 1947). I take this opportunity of expressing to Professor Copeland my hearty thanks for his unfailing kindness and courtesy in answering questions. As, however, the publication of Genera Filicum has ante-dated the publication of this paper, it has been possible to give page references and quotations that otherwise would have been postponed for a supplementary paper.

My thanks are due to Mrs. Allan (née Molesworth), until recently Botanist to the Auckland War Memorial Museum, for her very helpful assistance and co-operation, and also to others who have at different times kindly helped either by providing literature or by directing my attention to references. Even so, some precise references are sometimes omitted, as relevant literature is not available in this country. The names published below are listed in the order given in Genera Filicum. Numbers of heading refer either to the number of the family in the order, or the number of the genus in the family. [It should be noted, however, that a different classification has recently been published by R. E. Holtum, “A Revised Classification of Leptosporangiate Ferns” (Journ. Linn. Soc., Botany, October, 1947).]

Note: It has not been thought necessary to quote details of synonyms, etc., where these are readily obtainable in Cheeseman's Manual (1925).

Order Marattiales

4. Marattia Swartz

Marattia salicina Smith (1812).

Syn. Marattia fraxinea Smith (1790).

Christensen: Index Filicum and Supplements.

Order Filicales

3. Gleicheniaceae

This family has been revised by Christensen and Copeland. Christensen (Manual of Pteridology, 1938) divided its 120 odd species among five genera. Copeland (Philippine Journal of Science, Vol. 75, No. 4; 1941) followed Christensen, except that he carved out of Christensen's genus Sticherus Presl the small genus Hicriopteris, thus giving the family six genera.

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1. Gleichenia Smith

“Frond bipinnate or more compound, segments minute, round or ovate, sori usually composed of three or four sporangia. A small genus of the Oriental Tropics and farther south.” About ten species.

Copeland: Phillipine Journ. of Sci., Vol. 75, No. 4, 1941, p. 347.

Gleichenia microphylla (R. Br.) C. Christen

Syn. Gleichenia circinata Swartz (1801).

Carse: Trans. N.Z.I., Vol. 56, 1926, p. 83.

Gleichenia circinata (Sw.) C. Christen

Syn. Gleichenia dicarpa R. Br. (1810). Carse: Ibid., p. 83.

Gleichenia circinata var. hecistophylla Hooker f.

As regards the form G. circinata var. alpina, Carse remarks, “the form known as var. alpina, seems to pass by gradual steps from var. hecistophylla, some of the subalpine forms of which it is almost impossible to distinguish from alpina.”

Carse: Trans. N.Z.I., Vol. 56, p. 83.

4. Sticherus Presl

“Rhizome and frond paleaceous or glabrescent, frond in typical development pinnate or bipinnate, its subsequent division pseudo-dichotomous, the growth of the terminal bud being suppressed above each node; foliar segments borne on ultimate axes and usually on one of three preceding internodes, nodes without stipule-like appendages; veins once forked; sori one to six (most commonly four) sporangia. A large genus throughout the Oriental and American tropics.”

Copeland: Gleicheniaceae of New Guinea, Phil. Journ. Sci., 1941, p. 350.

Sticherus cunninghamii (Hew. ex Hook.) Ching

Syn. G. cunninghamii (Hew. ex Hook. 1844).

Ching: Sunyatsenia, Vol. 5, 201–228, 1940.

Sticherus flabellatus (R. Br.) H. St. John Syn. G. flabellata R. Br. (1810).

H. St. John: Occasional Papers of the Bishop Museum, Vol. 17, No. 7, p. 81.

5. Dicranopteris Bernhardi

“Rhizome and axis with hairs (or naked) but no scales. Frond pinnate or forked at top of stipe, then repeatedly forked with a pair of ‘stipules’ subtending the forking; only stipules and ultimate segments foliaceous. Veinlets more than once forked. Sori with six to twelve sporangia.”

Copeland remarks: “A small genus if D. linearis is broadly construed, a fairly large genus if its many forms are treated as species. Pantropic a little beyond the Tropic of Cancer and South to New Zealand.”

Copeland: Gleicheniaceae of New Guinea, Philippine Journal of Science, Vol. 75, No. 4, p. 349.

Dicranopteris linearis (Burm.) Underwood Syn. G. linearis C. G. Clarke (1880).

5. Hymenophyllaceae

Christensen divided this great family of over six hundred species into four genera, the monotypic endemic genus Cardiomanes, the

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monotypic genus Serpyllopsis, and the two great genera Hymenophyllum and Trichomanes, distinguished from each other according to the shape of the indusium and the length of the soral receptacle. Copeland, however, has shown these features to be of unsound diagnostic value; moreover, he finds, “The family is diversified beyond the suspicion of those who have not given it particular study.” He has, then, split the family into thirty-three genera, of which New Zealand possesses fourteen. It is to be noted that most of the “new names” are actually old. “I find satisfaction,” remarks Copeland, “in the fact that most of the genera I recognise are long since set up or maintained by Presl and Van Den Bosch. Presl must have had a remarkable eye for genera.” He mentions that the bad state of nomenclature of the group has been the subject of much comment, “sometimes by those who contributed to the evil!” and adds “I have tried to bring order into the generic nomenclature by a scrupulous observance of the accepted rules.”

Copeland: Genera Hymenophyllacearum, The Philippine Journal of Science, Vol. 67, No. 1, 1938.

1. Mecodium Presl

As regards this name, Copeland remarks, “Presl certainly did not adequately describe his genus Mecodium. His description would be inaccurate, even if what he wrote were correct, which it is not!… I adopt his generic name because it exists, and its use is preferable to the invention of a new name.”

The largest genus of the family, pantropic and austral, c. one hundred species, with thirteen in New Zealand. Margins of fronds smooth and naked (except three species not found in New Zealand). Some species have fronds less delicate than is usual with the family. Sorus simple.

Copeland: Genera Hymenophyllacearum, 1938, p. 17.

Mecodium sanguinolentum (Forster) Presl

Syn. Hymenophyllum sanguinolentum G. Forst. (Prod. 1786).


Copeland: Genera Hymenophyllacearum, 1938, p. 19.

Mecodium flabellatum (Labill.) Copeland

Syn. Hymenophyllum flabellatum Labill (1806).

New Zealand and Polynesia.

Copeland: Genera Hymenophyllacearum, 1938, p. 21.

Mecodium rufescens (Kirk) Copeland

Syn. Hymenophyllum rufescens Kirk (1879).


Copeland: Genera Hymenophyllacearum, 1938, p. 21.

This fern is very closely allied to M. flabellatum, intermediate forms are found between the two.

Holloway: Studies in New Zealand Hymenophyllaceae, Trans. N.Z.I., Vol. 54.

Mecodium rarum (R. Br.) Copeland

Syn. Hymenophyllum rarum. R. Brown (1810).

New Zealand and Western Australia.

Copeland: Genera Hymenophyllacearum, 1938, p. 21.

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Mecodium montanum (Kirk) Copeland

Syn. Hymenophyllum montanum Kirk (1877).


Copeland: Genera Hymenophyllacearum, 1938, p. 22.

“There is a single specimen, Kirk, 564, in the United States National Herbarium, topotype if not a cotype. To Kirk's description, it may be added that the internal walls are thickened and obscurely pitted. The receptacle is like that of H. rarum, to which it is more nearly related than to H. javanicum, with which Kirk compares it, but from which it differs in having narrow segments and decidedly ragged lips. New Zealand, apparently very rare.”

Copeland: Hymenophyllum, Philippine Journal of Science, Vol. 64, Nos. 1–2, 1937, p. 132.

Mecodium flexuosum (A. Cunn.) Copeland

Syn. Hymenophyllum flexuosum A. Cunningham (1836).

H. australe Willd., part auctt.


Copeland: Genera Hymenophyllacearum, 1938, p. 24.

“This is the New Zealand representative of the group H. badium [Note: in this species the receptacle commonly forks], not at all like that of H. javanicum, with which it has been confused strangely. It is recognisable at sight by the wide wing of the upper part of the stipe with crisped margin, but flat along the axis. Endemic in New Zealand, probably common in both islands.”

Copeland: Hymenophyllum, Philippine Journal of Science, Vol. 64, Nos. 1–2, p. 155.

Mecodium pulcherrimum (Colenso) Copeland

Syn. Hymenophyllum pulcherrimum Colenso (1844).


Copeland: Genera Hymenophyllacearum, 1938, p. 24.

Mecodium villosum (Colenso) Copeland

Syn. Hymenophyllum villosum Colenso (1844).


Copeland: Genera Hymenophyllacearum, 1938, p. 24.

This endemic species is very closely related to the widely distributed M. sanguinolentum; indeed, Dr. Holloway considers it possibly a derived mountain form of the latter.

Holloway: Ibid., (Trans., Vol. 54).

Mecodium australe (Willdenow) Copeland

Syn. Hymenophyllum australe Willdenow (1810).

New Zealand, Tasmania, Victoria.

Copeland: Genera Hymenophyllacearum, 1938, p. 24.

“I can see no near affinity between H. australe, which seems to me a reduced relative of H. demissum, and either H. crispatum or H. javanicum.”

Copeland: Hymenophyllum, Philippine Journal of Science. Vol. 64, Nos. 1–2, p. 159.

Mecodium atrovirens (Col.) Copeland

Syn. H. atrovirens Colenso (1844).


“Specimens of this species and M. australe sent me by… Prof. Holloway seem very distinct, M. atrovirens being smaller, narrower

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throughout, and much darker, with conspicuous two-cells-thick margin.”

Copeland: Notes on Hymenophyllaceae, Phil. Journ. of Science, Vol. 73, No. 4, p. 457.

Mecodium demissum (Forster) Copeland

Syn. Trichomanes demissum Forster (1786).

Hymenophyllum demissum (Forster) Swartz (1801).


Copeland: Genera Hymenophyllacearum, 1938, p. 24.

H. demissum (Forster). With the type material for H. aucklandicum in hand, I can distinguish it in no way from a small H. demissum; H. sanguinolentum is a local relative; H. australe a related dwarf.”

Copeland: Hymenophyllum, 1937, p. 160.

Mecodium dilatatum (Forster) Copeland

Syn. Trichomanes dilatatum Forster (1786).

Hymenophyllum dilatatum (Forster) Swartz (1801).


Copeland: Genera Hymenophyllacearum, Philippine Journal of Science, Vol. 67, No. 1, 1938, p. 24.

Mecodium scabrum (A. Rich) Copeland

Syn. Hymenophyllum scabrum A. Richard (1832).


Copeland: Genera Hymenophyllacearum, 1938, p. 24.

2. Craspedophyllum (Presl) Copeland

Craspedophyllum (Presl) Copeland

Copeland considers this genus is probably related to Mecodium. It is characterised by minute fronds, and the black contents of the obliquely placed margin cells with internal walls wavy thickened.

Copeland: Genera Hymenophyllacearum, 1938, p. 27.

Craspedophyllum armstrongii (Baker) Rae

Syn. Hymenophyllum cheesemanii Baker (1873).

Hymenophyllum cheesemanii var. armstrongii Cheeseman (1906).

Trichomanes armstrongii Baker (1868).

Copeland: Genera Filicum, p. 33.

Holloway was convinced that Hymenophyllum cheesemanii and H. cheesemanii var. armstrongii were simply forms of the same species, since the dark margin of var. armstrongii, which constituted its distinguishing feature, was not a stable character. (Trans. N.Z. Inst., 54, 1923, p. 592.) Copeland has now accepted this reduction.

4. Sphaerocionium Presl

A large tropical genus, mainly South American. The sorus is simple. It is distinguished from Mecodium by its pubescence, and from Leptocionium by the included receptacle. “There is closer apparent affinity to Microtrichomanes; if this be real, the latter may be regarded as a daughter genus… Characterised with emphasis on the pubescence, it is a natural genus, typified by S. hirsutum, which must be regarded as the type of Presl's genus… It should be possible to break so large a genus into natural groups,… but I do not know these plants intimately enough to do this with any confidence to

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the genus as a whole.”

Copeland: Genera Hymenophyllacearum, 1938, p. 29.

Sphaerocionium ferrugineum (Colla) Copeland

Syn. Hymenophyllum ferrugineum Colla (1836).

Hymenophyllum subtilissimum Kze. (1837).

Chile to New Zealand.

Copeland: Genera Hymenophyllacearum, 1938, p. 33.

Sphaerocionium lyallii (Hook. f.) Copeland

Syn. Hymenophyllum lyallii Hook. f. (1854).

Trichomanes lyallii Hook. f. (Syn. Fil., 1867).

New Zealand and New South Wales.

Copeland: Genera Hymenophyllacearum, 1938, p. 33.

Commenting on this species in the Manual, Cheeseman remarks: “Exactly intermediate between Trichomanes and Hymenophyllum as fas as the structure of the indusium is concerned. It is purely a matter of taste as to which genus it should be referred” (page 17). When classification becomes a matter of taste, revision is surely indicated!

5. Apteropteris Copeland

Copeland regards this montypic genus as a daughter genus of Sphaerocionium, but gives it generic status on account of the remarkable peculiarities of the frond. “The stellate hairs are unmistakable evidence of affinity to Sphaerocionium.”

Copeland: Genera Hymenophyllacearum, 1938, p. 35.

Apteropteris malingii (Hooker) Copeland

Syn. Trichomanes malingii Hooker (1862).

Hymenophyllum malingii (Hooker) Mettenius (1864).


Copeland: Genera Hymenophyllacearum, 1938, p. 35.

7. Hymenophyllum Smith

Closely allied to Meringium, from which it differs in the more deeply cleft involucre, the shorter receptacle, the absence of peculiarly pitted, thickened cell walls, and the unusually small size. Consists of about twenty-five very similar species, notable for the occurrence of some in temperate lands.

Copeland: Genera Hymenophyllacearum, 1938, p. 38.

Hymenophyllum revolutum Colenso

Syn. Hymenophyllum revolutum Colenso, Tasm. Journal, 2, 1844, 186.

Hymenophyllum zeelandicum Van Den Bosch, Ned. Kruid. Arch., 53, 1863, 175.

Hymenophyllum pusillum Colenso, Trans. N.Z. Inst., 12, 1880, 365.

?Hymenophyllum tunbridgense C. Chr. part., Cheeseman, Holloway, Trans. N.Z. Inst., 54, 1923, 592, pl. 68, viz. Smith.


“All of the ‘species’ treated here have been reduced to H. tunbridgense (or H. peltatum, in error), which I do not find present from New Zealand. For one thing, all New Zealand specimens have the rachis largely wingless.

“None of these has a frond five inches long, the range being from 8 cm. exclusive of stipe, down to less than 2 cm. (fertile). The receptacle is somewhat extruded, or equals the valves, or is shorter; I am quite sure that in this case the difference is not diagnostic. In poorly

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fruiting material the sori are axillary, as in H. tunbridgense, but on contracted segments. In fuller fructification the sori tend to become terminal.

“This is as I see it. But I know well that the man in the field can know his plants better than the man in the herbarium, and on that ground would mistrust my judgment in reducing Colenso's three species to two or one, except that the reduction is accepted by Holloway. However, dwarfs in general are derived from plants of more usual size. The two being surely nearly related, H. pusillum would probably be derived from H. revolutum, and could hardly become specifically fixed in the actual presence of the ancestral form; and they have the same type locality—Lake Waikare. If they are distinct, the name of the smaller form is H. zeelandicum.

“I have no specimen fitting the description of H. pygmaeum, which is very like that of H. minimum. Colenso, however, was very positive that they are distinct and not nearly related.”

Copeland: Hymenophyllum, 1937, p. 89.

8. Meringium Presl

Closely allied to Hymenophyllum, but lower part of the indusium closed and the receptacle projecting some distance from the sorus. Distinguished in general by larger coarser fronds with thick and coarsely pitted walls. New Zealand species less distinct from Hymenophyllum in development of the walls.

Copeland: Genera Hymenophyllacearum, 1938, p. 39.

Meringium minimum (A. Rich.) Copeland

Syn. Hymenophyllum minimum A. Rich (1832).

Copeland: Notes on Hymenophyllaceae, Philippine Journal of Science, Vol. 73, No. 4, p. 457.

“Excellent material from Professor Holloway shows the receptacle so constantly and so far protruding that the propriety of treating the dwarf as a Meringium is evident. It may well be remembered, not only that distinctions may become inconspicuous with dwarfing, but that Meringium in its full and typical development is a New Guinea-Malayan genus, and that in New Zealand and America, nearer to the common ancestral source, it is always less distinct from Hymenophyllum.”

Copeland: Notes on Hymenophyllaceae, p. 437.

Meringium multifidum (Forster) Copeland

Syn. Trichomanes multifidum Forster (1786).

Hymenophyllum multifidum (Forster) Swartz (1801).


Copeland: Genera Hymenophyllacearum, 1938, p. 44.

Meringium bivalve (Forster) Copeland

Syn. Trichomanes bivalve Forster (1786).

Hymenophyllum bivalve (Forster) Swartz (1801).

New Zealand to Queensland.

Copeland: Genera Hymenophyllacearum, 1938, p. 44.

16. Cardiomanes Presl

A monotypic endemic genus, perhaps the most isolated in the whole family.

Copeland: Genera Hymenophyllacearum, 1938, p. 50.

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Cardiomanes reniforme (Forster) Presl

Syn. Trichomanes reniforme Forster (1768).

Copeland: Trichomanes, Philippine Journ. of Science, Vol. 51, No. 2, 1933, p. 270.

17. Vandenboschia Copeland

Copeland described this genus as, “The most nearly cosmopolitan genus in the family. Among those which have been included in Trichomanes, it is the least differentiated; on this ground I regard is as the most nearly primitive. A considerable number of the genera of more limited geographic range are evidently derived from it.” Indusium funnel-shaped, receptacle thin and protruding.

Copeland: Genera Hymenophyllacearum, 1938, p. 61.

Vandenboschia colensoi (Hooker f.) Copeland

Syn. Trichomanes colensoi Hooker f. (1854).


Copeland: Genera Hymenophyllacearum, 1938, p. 52.

18. Polyphlebium Copeland

Phlebiophyllum Van Den Bosch, Versl. Akad. Wet. Amsterdam, 11, 1861, 321, non Phlebophyllum Ness (1832).

An isolated genus of only one species confined to New Zealand, Tasmania and Queensland. “The only plant in the family with very thin leaves in which the veins branch freely in undivided segments of the frond.”

Copeland: Genera Hymenophyllacearum, 1938, p. 55.

Polyphlebium venosum (R. Br.) Copeland

Syn. Trichomanes venosum R. Br. (1810).

New Zealand to Tasmania and Queensland.

Copeland: Genera Hymenophyllacearum 1938, p. 55.

21. Crepidopteris Copeland

A small group of Malayan and Polynesian origin, with small bi-pinnate fronds. Indusium funnel-shaped, receptacle protruding. Copeland remarks: “The longitudinal elongate marginal cells are a convenient diagnostic characteristic, but the genus is recognisable at sight by a combination of characters—small size, thinness, narrowly elliptic form, degree of dissection, position of sori—which are not individually diagnostic. If two rows of cells are elongate, the inner row may be double in thickness.”

Copeland: Genera Hymenophyllacearum, 1938, p. 57.

Crepidopteris endlicheriana (Presl) Copeland

Syn. Trichomanes endlicherianum Presl (1849–1851).


Copeland: Genera Hymenophyllacearum, 1938, p. 58.

This species has been known previously in New Zealand as Trichomanes humile (now Crepidopteris humilis). Actually C. humilis is not found in New Zealand. C. endlicherianum differs from it as follows. It has two rows of marginal cells instead of one, and these cells are more elongate, the frond is narrower, though its segments are actually longer, and the wing of the involucre is wider so that the mouth does not project beyond it as much as in C. humilis. The receptacle, apparently less fragile, is longer.

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Copeland: Trichomanes, Philippine Journal of Science, 1933, p. 168.

31. Selenodesmium Copeland

A well-defined pantropic group of ill-defined species. The name probably refers to the semilunar cross-section of the bundle of the stipe.

Copeland: Genera Hymenophyllacearum, 1938, p. 80.

Trichomanes rigidum Sw. type species.

A small, ground-loving genus of about ten species. Fronds typically triangular. Indusium cylindrical with very long protruding receptacle. Reaches “more typical development in Malay-Polynesian region. Plants strictly terrestrial and intolerant of desiccation.” (Stout prop-roots usually support stems of old plants.)

Copeland: Genera Hymenophyllacearum, 1938, p. 80.

Selenodesmium elongatum (A. Cunn.) Copeland

Syn. Trichomanes elongatum A. Cunn. (1836). New Zealand and Eastern Australia. Copeland: Genera Hymenophyllacearum, 1938, p. 82.

33. Macroglena Copeland

Genus of twelve species found in tropics of Old World, and south to New Zealand and Madagascar.

“The extremely narrow (or eventually wanting lamina), which is the conspicuous characteristic of this genus, is obviously not a primitive character, but may with confidence be regarded as the result of reduction…. As I construe the genus, it includes T. strictum of New Zealand, and is old enough to be of direct Antarctic origin. Macroglena as a genus is unspecialised either as to the length of the rhizome or the structure of the cell walls—items usually characteristic of genera in this family.”

Copeland: Genera Hymenophyllacearum, 1938, p. 83.

Macroglena stricta (Menzies) Copeland

Syn. Trichomanes strictum Menzies (1829). Endemic.

Copeland: Genera Hymenophyllacearum, 1938, p. 85. Note: Hymenophyllum peltatum is the only member of the family whose name remains unchanged. It is, however, now included under Hymenophyllum Smith (about twenty-five species) as interpreted above, not the genus Hymenophyllum Smith as interpreted by Christensen (Verdoorn, Manual of Pteridology, p. 531), which includes about 300 species.

6. Pteridaceae

23. Hypolepis Bernhardi

Hypolepis rugosula (Labill.) J. Smith

Syn. Dryopteris punctata (Thunberg) C. Christen. (1905). This species has had a troubled history. Both H. rugosula and H. punctata were united by Hooker and Baker under Polypodium punctata (Syn. Fil., p. 312, 1873). They are listed in the Manual as Dryopteris punctata. Christensen, however, in correspondence with Carse, states: “The genus Hypolepis has no true ‘scales’; the investiture is always… by hairs…. By this character Hypolepis can always be certainly known from Dryopteris,”

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H. rugosula is distinguished from H. punctata by its red rachises and its viscosity.

Carse: Trans. N.Z.I., Vol. 56, 1926, p. 82.

Hypolepis punctata (Thunb.) Mett. in Kuhn, Fil. Ar., 1868, 120. Syn. H. petriana Carse (1918).

This species was for a long time thought to be a form of H. tenuifolium. Then Carse described it as an independent endemic species under the name H. petriana (Trans. N.Z.I., Vol. 50, 1918, p. 64); however, when he subsequently submitted specimens to the late Dr. Christensen, that authority declared them to be H. punctata. H. punctata differs from both H. tenuifolia and H. rugosula in its pale almost naked stipes and rachises.

Carse; Ibid., 1926, p. 81.

28. Pteridium Gleditsch

Pteridium esculentum (Forst.) Diels (1899).

Syn. Pteridium aquilinum var. esculentum (Forst.) Kuhn (1882).

Cockayne: Report on a Botanical Survey of the Tongariro National Park, Department of Lands Report, p. 34, 1908.

Note: R. M. Tryon, jun., in Revision of the Genus Pteridium, considers the genus consists of one species, Pteridium aquilinum (L.) Kuhn, and eleven varieties, our plant being Pteridium aquilinum var. esculentum. Copeland, however (Genera, p. 60), thinks the forms “better treated as six or more species, even though they blend in places.”

31. Pteris L.

Pteris saxatilis Carse

Syn. P. macilenta A. Rich. var. saxatilis Carse (1919).

This change was made by Carse, who remarks: “After some years further study of this plant, I am convinced that it is very distinct from P. macilenta (Trans. N.Z.I., Vol. 59, 1928, p. 315).

39. Cheilanthes Swartz

Cheilanthes distans (R. Br.) Mett.

Syn. Nothoclaena distans R. Br. (1810).

The genera Cheilanthes and Notholaena are very closely allied. Notholaena was reduced to Cheilanthes by Domin in 1915 (Bibl. Bot., 85, 133). The reduction is accepted by Copeland in Genera Filicum. He states “N. distans is by affinity a Cheilanthes in the strictest reasonable sense” (p. 66).

It was described by Field in New Zealand Ferns (1890) as Cheilanthes n.sp. (Mettenius never accepted the genus Notholaena.)

60. Anogramma Link

Small ferns with very small and nearly naked rhizomes. Fronds tufted, subdimorphous, two to three pinnate. Veins free. Sporophyte ephemeral (annual?). Prothallus lobed and perennial with tuber-like archegoniophores, which may propagate vegetatively.

Christensen: Man. of Pteridology, p. 538).

Anogramma leptophylla (L.) Link (1841).

Sya. Gymnogramme leptophylla Desv. (1811).

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11. Cyathaceae

3. Cyathea Smith

This genus was originally divided into three genera, Cyathea, Alsophila and Hemitelia. The genera were characterised by the indusia, which was described as “globose” in Cyathea, “never covering the sorus, … usually a half cup-shaped or semi-circular scale on the lower side of the sorus” in Hemitelia, and as “altogether wanting” in Alsophila (Cheeseman, Manual, p. 22, 25, 26). Further research has not justified the use of these apparently straightforward distinctions. Copeland united the three genera into one in 1909 (Philippine Journal of Science, Bot. 4, 28 ff.). Writing in 1929, Copeland remarks: “Alsophila and Hemitelia, as commonly construed, are not genera in any modern scientific sense. Alsophila is a catch-all for species without an indusium and specimens so collected; Hemitelia for those with the indusium incomplete. But the tree ferns are not divisible naturally into any two or three or more groups with indusia characters.” (The Oriental Genera of Polypodiaceae, University of California, Publications in Botany, Vol. 16, 1929, p. 54.) Domin follows him, remarking, “there is hardly any living pteridologist who would not share the opinion that the three genera are untenable and quite unnatural units, connected by many transitions.” (The Species of the Genus Cyathea: Acta Botanica Bohemica, Vol. 9, 1930, p. 85). Domin also includes in Cyathea Bower's family Protocyathaceae. As constituted by Domin the genus includes about 770 species (800 according to Copeland, 1947).

Cyathea smithii (Hook. f.) Domin

Syn. Hemitelia smithii Hooker f. (1865).

Domin: Pteridophyta, 1929, p. 264.

Cyathea colensoi (Hook. f.) Domin

Syn. Alsophila colensoi Hook. f. (1864).

Domin: Ibid., p. 262.

Cyathea falciloba (Col.) Domin

Syn. Hemitelia falciloba Col., T.N.Z.I., 24, 1892, 394.

Domin: Ibid., p. 264.

Cheeseman lists this fern in the Manual as (?) Hemitelia falcifolia and gives it as a synonym of Cyathea dealbata. Colenso describes it as Hemitelia falciloba and says the indusium is “a very small concave greyish scale on the posterior side, margin entire.” He stated he discovered two specimens of H. falciloba “on the side of a precipitous gully overhanging a small streamlet in a forest south of Dannevirke, January, 1889.” Unfortunately, he never discovered any more, and on his next visit he found the original two had been burnt. The plant has never been recorded since.

Cyathea novae-zelandiae (Col.) Domin

Syn. Hemitelia smithii var. microphylla Cheeseman (1906).

Hemitelia microphylla Col. (1895). Domin: Ibid., p. 264.

Originally described by Colenso under Hemitelia microphylla (Trans. N.Z.I., Vol. 27, 1895, p. 399). Later it was reduced to varietal rank. Domin has again made it an independent species. As, however,

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Cyathea microphylla Mett., a South American fern, was described in 1856, it had priority; consequently it was necessary to find another name for the New Zealand fern.

12. Aspidiaeae

10. Polystichum Roth

Polystichum mohrioides (Bory) Pr. Tent. 83, 1836.

This fern, which was described by Kirk (Trans. N.Z.I., 14, 1882, p. 386) from a form “collected on the Auckland Islands by a sailor, during the past year.” Cheeseman omitted it from the Manual, as he did not think it grew in New Zealand. One plant, however, has now been found growing in dry Leptospermum scrub on Kapiti Island.

Allan: Trans. N.Z.I., 65, 1936, p. 231.

16. Rumohra Raddi

Terrestrial ferns of moderate size with (usually) long creeping rhizomes, entire or subentire paleae, and lamina anadromic in plan. Ultimate pinnules usually rhomboid and aristate. Veins free. Indusium peltate in the type (R. adiantiformis) but orbicular-reniform in most species. About fifty species. Copeland, also Ching (Sinensia, 5, 1934, 33) include in Rumohra the genus Polystichopsis (Genera, p. 113). Holtum does not accept the reduction (Journ. Linn. Soc. Botany, Oct., 1947, p. 152).

Rumohra adiantiformis (Forst.) Ching

Syn. Polyatichum adiantiforme J. Smith (1875).

Ching: Sinensia, 5, 23, 1934, p. 5. This fern is the type species of Rumohra.

Rumohra aristata (Forst.) Ching

Syn. P. aristatum (Forst.) Presl (1836). Ching: Ibid.

Rumohra hispida (Sw.) Copeland

Syn. Polystichum hispidum (Sw.) J. Smith (1841). Copeland: Genera Filicum, p. 114.

32. Ctenitis Christensen

Terrestrial ferns, moderate or larger, rhizome usually short, fronds bi-pinnatifid, axes usually with toothed paleae, typically hairy above with articulate pluricellular hairs, costae not decurrent, veins free. Sori with indusium round reniform or wanting. Christensen estimated about 150 species. Copeland includes in the genus also Parapolystichum. Before making his new combinations, Copeland remarks, “As it has been Christensen's service to recognise this genus as a natural one, and thereby to contribute more than in any other single respect to the proper classification of the great aggregate of species known of late as Dryopteris, I would gladly have left it to him to make the combinations he indicated.”

Copeland: Genera, p. 123.

Ctenitis glabella (A. Cunn.) Copeland

Syn. G. glabella (A. Cunn.) C. Christensen (1905). Copeland: Genera Filicum, p. 124.

Ctenitis decomposita (R. Br.) Copeland Syn.

D. decomposita (R. Br.) Ktze. (1891). Copeland: Genera Filicum, p. 124.

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Ctenitis velutina (A. Rich.) Copeland

Syn. D. velutina (A. Rich.) O. Ktze. (1891). Copeland: Genera Filicum, p. 124.

51. Thelypteris Schmidel

Terrestrial ferns of moderate size, paleae rarely dense, often pubescent, frond typically pinnatifid, usually narrowed towards both ends, often hairy with simple unicellular hairs. Veins free. Sori small, round (rarely elongate), indusium when present round-reniform. About 500 species, mostly of Europe and other north temperate lands.

This description is taken from Copeland (Genera, p. 136). Copeland, however, adopts the name Lastrea for the genus, as he regards Schmidel's publication of Thelypteris as invalid. Alston, however, has vigorously taken up the cudgels on behalf of Thelypteris (Kew Bulletin, 1932, p. 309), and the present tendency among British botanists appears to be to adopt Thelypteris.

Thelypteris palustris Schott var. squamigera (Schlecht)

Syn. Dryopteris thelypteris Asa Gray var. squamulosum Schlecht.

Aspidium squamigerum (Schlecht) Adumb. 23, pl. 11 (1825).

Weatherby: Contributions from the Gray Herbarium of Harvard University, 1924, p. 40.

Originally thought to be confined to New Zealand and South Africa, var. squamigera was also collected in Portuguese West Africa by Mrs. R. C. Curtis. It was determined by Mr. Weatherby, of the Gray Herbarium, who published the name cited above as a new combination. I am indebted to Miss L. B. Moore for directing my attention to this reference.

Thelypteris uliginosa (Kze.) Ching

Syn. Dryopteris setigera (Blume) O. Ktze.

R. E. Ching: Bull. Fan Memorial Institute, Botany, Vol. 6, Mar., 1936, p. 345.

In the same Bulletin, Ching also made the combination Thelypteris setigera (Blume) Ching, but he states that this species does not grow beyond tropical Asia. The Polynesian and Australian plant known as Dryopteris setigera (Blume) is a different species altogether. In making the combination Thelypteris uliginosa he comments, “the nomenclatural confusion for this widely distributed but constant fern is almost unthinkable.” Anyone who has tried to disentangle the synonymy will agree!

52. Cyclosorus Link

Cyclosorus is so intimately related to Thelypteris that Holtum (Journ. Linn. Society London, Botany, Vol. 53, p. 130) thinks them better combined. Copeland, however, thinks it more convenient to keep them separate. The distinguishing characteristic is that in Cyclosorus the veins in adjacent segments characteristically meet and unite at or below the segments. A genus of probably 300 species.

Christensen: Manual, p. 545 and Copeland: Genera, p. 141.

Cyclosorus gongylodes (Schkuhr) Link Syn. Aspidium gongylodes Schkuhr (1809). Ching: Bull, Fan, 8, 1938, 235?.

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Cyclosorus pennigerus (Forst.) Copeland

Syn. Dryopteris pennigera (Forst.) C. Christensen (1905). Copeland: Genera, p. 143.

Cyclosorus dentatus (Forst.) Ching

Syn. Dryopteris parasitica Kuntze (1891).

There is a double revision here. Dryopteris parasitica was revised by Carse as D. dentatus (Trans., Vol. 56, 1926, p. 81) since “the typical parasitica of Linnaeus is Chinese, and. inter alia characterised by the frond being narrowed downwards.” (C. Chr.) Later the generic name was revised by Ching, who deals with the genus Cyclosorus in Bull. Fan., 8, 1938, 235, and presumably made the change there.

62. Athyrium Roth

Ground-loving ferns with short creeping to erect rhizomes. Both these and stipe bases clothed with chaffy scales. Sori elongate along the veins, indusium typically bent across the vein at the upper end of the sorus, or on both sides of the vein, making the sorus double. In a few species the indusium is wanting. The identity of Diplazium and Athyrium was first shown by Milde. Copeland remarks that the supposed distinction between the two genera is that Diplazium “has more or less symmetrical double sori occupying both sides of the fertile veinlets; the latter (Athyrium) sori bent across the veinlet, continuous from one side to the other at the upper end and two sides usually unequal… But in one after another of the natural groups of species there occur sori of both types, so that natural genera, defined by these characteristics do not exist.” (The Oriental Genera of Polypodiaceae, Univ. Calif. Publications in Botany, 1929, p. 69).

Athyrium australe (R. Br.) Presl, 98, 1836.

Syn. Athyrium umbrosum (Ait.) Presl (1836).

In his Ind. Fil., Suppl. 3, Christensen split up the A. umbrosum of Ind. Fil. (1906) into five species: A. umbrosum (Ait) Pr. (Atlantic Islands), A. australe (R. Br.) Pr. (Australia, New Zealand, New Caledonia), A. pseudosetigerum Christ. (Central Asia), A. bellum (Clarke) Ching (India), A. moricatum (Mett.) C. Chr. (Java).

Athyrium japonicum (Thunberg) Copeland

Syn. Diplazium japonicum (Thunberg) Bedd. (1876).

13. Blechnachae

1. Blechnum L.

Blechnum procerum (Forst. f.) Labill. Syn. B. capense Schlect (1825).

J. C. Andersen: Popular Names of New Zealand Plants (Trans. N.Z.I., Vol. 56, 1926, p. 681).

This would appear to be a compound species. One of its forms has the lower pinnae (almost invariably) not narrowed at the base, and stipes very long in proportion to the frond. There is no doubt that the species needs revision, and in the present state of our knowledge it seems impossible to come to final conclusions about some of its forms. Carse submitted characteristic B. capense (as then understood in New Zealand) to Dr. Christensen, who said that it “should probably be named B. procerum”

Carse: Trans. N.Z.I., Vol. 56, 1926, p. 82.

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Blechnum minor (R. Br.) Ckne.

Syn. B. capense var. minor Hook f. (1855).

Blechnum patersoni Mett. (1856)

Syn. Blechnum patersoni var. elongatum Hook. and Bak. 1873.

The type and its variety were originally thought two forms. There was, Cheeseman remarks, “The type which was believed to always have simple fronds, and secondly var. elongatum in which the fronds were pinnatifid…. But it was long since proved that the variety produces simple fronds and B. patersoni has been found with pinnatifid ones, so that the supposed distinction falls to the ground.”

Cheeseman: The Subantarctic Islands of New Zealand, Vol. 2, 1909, p. 439.

14. Aspleniaceae.

1. Asplenium L.

Asplenium lamprophyllum Carse

This fern was long thought to be a form of A. bulbiferum, from which it is distinguished by the narrow linear sorus, bright shining leaf surface, and absence of bulbils.

North Island as far south as Te Aroha. Carse: Trans. N.Z.I., Vol. 56, 1926, p. 81.

Asplenium falcatum Lam.

Syn. Asplenium adiantioides (Lin.) C. Christen. (1905).

The name Asplenium adiantioides (Lam.) is now applied to A. praemorsum Swartz. Copeland remarks, “There is no choice but to adopt Lamark's name for this species, letting A. adiantioides (Linnaeus) C. Christensen be known again by its long-familiar name, A. falcatum.”

Copeland: Pteridophytes of the Society Islands, Bishop Museum Bull., 93, p. 49.

Asplenium colensoi Colenso

Syn. A. hookerianum var. colensoi Moore (1858).

This fern was originally named and described as an independent species by Colenso. Later it was reduced to a variety of hookerianum, with which it is connected by intermediate forms; indeed, Cheeseman says, “occasionally fronds of both forms can be found on the same plant” (Manual, p. 50). Cockayne and Allan, however, reinstated it as a species, the intermediate forms being described as hybrids (Cockayne and Allan: An Annotated List of Wild Hybrids found in New Zealand, Annals of Botany, Jan., 1934). Described in the Tasmanian Journal of Natural Science, Vol. 2, 1845, p. 170. (I am indebted to Miss Molesworth for this reference.)

5. Pleurosorus Fée

A small genus of about three species. It is related to Asplenium, but lacks an indusium and is clothed with articulated hairs.

Christensen: Man. of Pteridology, p. 541.

Pleurosorus rutifolius (R. Br.) Fée

Syn. Gymnogramme rutaefolia Desv. (1827).

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16. Polypodiaceae

25. Pyrrosia Mirbel

A genus of about 100 species of rock-dwelling or epiphytic ferns, rhizomes densely scaly and articulate at base. Fronds simple, usually entire, rarely hastately or palmately lobed, texture generally leathery, sori rounded, rarely oblong, exindusiate. A striking characteristic of the genus is the presence of stellate hairs on the frond, usually persistent on the underside, and brown to white in colour, giving the genus its name (purros, flame-coloured).

Ching: Bull. Chinese Bot. Soc., Vol. 1, No. 1, 1935, pp. 36–72.

Pyrrosia serpens (Forst.) Ching

Syn. Pyrrosia serpensis (Forst.) Ching. Bull. Chin. Bot. Soc., 1, 1935, p. 49. Pyrrosia serpens, published by Oliver, Dominion Museum Records.

vol. 1, Oct., 1944, p. 221.

Cyclophorus serpens, (Forst.) C. Christen, (1905). Clung: Bull. Chin. Bot. Soc., 1, 1935, p. 49.

28. Microsorium Link

Microsorium was revived by Copeland (Univ. Calif. Publ., 16, 1929, 111). Christensen in the Index Filicum (1934) reduced it to Polypodium; however, he recognised Microsorium as a valid genus in the Manual of Pteridology (1938). As the name suggests, this genus is characterised by very small sori. These are punctiform and numerous. Stipes articulate, fronds simple, rarely pinnatifid, and usually thin in texture with distinct reticulate venation. Copeland includes here Phymatodes Presl. Christensen and Ching keep this genus separate. Ching recognises about 40 species.

Copeland: Genera, p. 193.

Microsorium diversifolium (Willd.) Copeland

Syn. Polypodium diversifolium Willd. (1810).

Copeland: Genera Filicum, p. 196.

Microsorium novae-zealandiae (Baker) Copeland

Syn. Polypodium novae-zealandiae Baker (1886).

Copeland: Genera Filicum, p. 196.

Microsorium pustulatum (Forst.) Copeland

Syn. Polypodium pustulatum Forst. (1786).

Copeland: Genera Filicum, p. 196.

52. Grammitis Swartz

A genus of about 150 species. Small or minute plants, usually epiphytic. Rhizome paleate, usually short, stipes non-articulate, fronds simple, linear or lanceolate, rarely shallowly lobed, costae usually prominent, veins typically free. Sori typically forming single rows each side of the costae, sometimes impressed or immersed, exindusiate.

Copeland: Genera, p. 210.

Copeland emphasises the fact that Grammitis, Xiphorpters and Ctenopteris (and sometimes Cheiropleuria) “excepting the Matoniaceae themselves are the only members of the phylum with globosetetrahedral spores without epispore” (Genera, p. 221). Copeland prepared a monograph on the genus based on material of almost all species. It was sent to Manila before the outbreak of war and its fate is not known. He also made diagnoses of 26 new species in Univ. Calif. Publ. Bot., 18, 1942, 222.

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Grammitis billardieri Willd. (1810).

Syn. Polypodium australe Mett. (1857).

Grammitis ciliata Col. (1844).

Syn. Polypodium australe var. villosum Hooker f. (1855).

Grammitis pumila Armstrong (1882)

Syn. Polypodium australe var. pumilum Cheessman (1906).

Grammitis crassa Fée (1850–52).

Syn. Polypodium billardieri (Willd.) C. Christensen (1906).

This species is found in the Auckland Islands. It is similar to G. billardieri, but much larger and the fronds are much thicker.

62. Anarthropteris Copeland

This is a new genus created by Copeland, who considers it most closely related to Loxogramme, from which it is distinguished by the soral paraphyses and “in general by the position and shape of the sori and by the spores.” Anarthropteris dictyopteris has, in the past, frequently been classified with ferns whose stipes are articulated to the rhizome. Copeland underlines the absence of articulation by naming his new genus Anarthropteris (unjointed).

Copeland: Genera, p. 218.

Anarthropteris dictyopteris (Mett.) Copeland

Syn. Polypodium dictyopteris Mett. (1861).

Copeland: Genera Filicum, p. 218.

63. Ctenopteris Blume

A large genus of probably more than 200 species. It is most closely related to Grammitis; indeed, Ching combined the two genera. Copeland, however, regards the resulting genus as “quite too inconvenient” and retains Ctenopteris. It has been included in the genus Polypodium by most authors including Christensen. Differences are in regard to spores, typical lack of articulation in Ctenopteris, which also has fronds hairy or glabrescent and always on a contracted base, while those of Polypodium are paleate or naked, usually on a broad base. The veins in Ctenopteris are almost always simple; they fork at least once in Polypodium. The sporangial pedicel of Ctenopteris has usually one row of cells, while that of Polypodium has three.

Copeland: Genera, pp. 218–219.

63. Ctenopteris Blume

Ctenopteris grammitidis (R. Br.) J. Smith. (1875).

Syn. Polypodium grammitidis R. Br. (1810).

16. Salviniaceae
2. Azolla Linn.

Azolla filiculoides var. rubra (R. Br.) Diels

Syn. Azolla rubra R. Br. (1810).