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NEW SOUTH WALES 

DEPARTMENT OF AGRICULTURE 


CONTRIBUTIONS 

FROM THE 

NEW SOUTH WALES 

NATIONAL HERBARIUM 



Flora Series: No. 201 


1963 


a 99777 













CONSPECTUS OF FAMILIES OF VASCULAR PLANTS 
REPRESENTED IN THE FLORA OF NEW SOUTH WALES 

(The account of each family, or consecutive group of families as issued, will appear with 
separate pagination; the names of those families that have been published to date are in italics.) 


GYMNOSPERMAE 


1 

Zamiaceae 

4 

Pinaceae 

2 

Podocarpaceae 

5 

Cupressaceae 

3 

Araucariaceae 




ANGIOSPERMAE- 

-MONOCOTYLEDONI 

6 

Typhaceae 

28 

Eriocaulaceae 

7 

Sparganiaceae 

29 

Commelinaceae 

8 

Pandanaceae 

30 

Pontederiaceae 

9 

Potamogetonaceae 

31 

Philydraceae 

10 

Ruppiaceae 

32 

Juncaceae 

11 

Zannichetliaceae 

33 

Liliaceae 

12 

Najadaceae 

34 

Xanthorrhoeaceae 

13 

Zosteraceae 

35 

Agavaceae 

14 

Aponogetonaceae 

36 

Philesiaceae 

15 

Posidoniaceae 

37 

Smilacaceae 

16 

Juncaginaceae 

38 

Haemodoraceae 

17 

Alismataceae 

39 

Amaryllidaceae 

18 

Hydrocharitaceae 

40 

Hypoxidaccae 

19 

Gramineae, Part l 

41 

Alstroemeriaceae 

20 

Cyperaceae 

42 

Dioscoreaceae 

21 

Paimae 

43 

Petermanniaceae 

22 

Araceae 

44 

Iridaceae 

23 

Lemnaceae 

45 

Zingiberaceac 

24 

Flagellariaceae 

46 

Cannaceae 

25 

Restionaceae 

47 

Burmanniaccae 

26 

Centrolepidaceae 

48 

Orchidaceae. See 

27 

Xyridaceae 


Orchids of New 5 


ANGIOSPERMAE—DICOTYLEDONES 


49 Casuarinaceae 

50 Piperaceae 

51 Salicaceae 

52 Fagaceae 

53 Ulmaceae 

54 Moraceae 

55 Cannabiaceae 

56 Urticaceae 

57 Proteaceae 

58 Loranthaceae 

59 Santalaceac 

60 Olacaceae 

61 Aristolochiaceae 

62 Polygonaccae 

63 Chenopodiaccae 

64 Dysphaniaceae 

65 Amaranthaceae 

66 Nyctaginaceae 

67 Phytolaccaceae 

68 Gyrostemonaceae 

69 Molluginaceae 

70 Aizoaceae 


71 Portulacaceae 

72 Basellaccae 

73 Caryophyllaceae 

74 Nymphaeaceae 

75 Cabombaceae 

76 Ceratophyllaceae 

77 Ranunculaceae 

78 Berberidaceae 

79 Menispermaceae 

80 Winteraceae 

81 Annonaceae 

82 Eupomatiaceae 

83 Trimeniaceae 

84 Moniraiaceae 

85 Lauraceae 

86 Cassythaceae 

87 Papaveraceae 

88 Fumariaceae 

89 Crucifcrae 

90 Capparidaceae 

91 Resedaceae 

92 Droseraceae 


(Continued on inside back cover) 


NEW SOUTH WALES 


DEPARTMENT OF AGRICULTURE 


CONTRIBUTIONS 

FROM THE 

NEW SOUTH WALES 

NATIONAL HERBARIUM 



Flora Series: No. 201 1963 

Sydney: V. C. N. Blight, Government Printer—1963 


G 99777—1 












/ 


FLORA OF NEW SOUTH WALES 


by 

VARIOUS BOTANISTS 


Produced under the Direction of 

R. H. ANDERSON 

Director and Chief Botanist, 
National Herbarium of New South Wales 


Pteridophyta of South Eastern Australia 

by 


MARY D. TINDALE 


No. 201 HYMENOPHYLLACEAE 


201. Hymenophyllaceae. ( Tindale) 


1 


201. HYMENOPHYLLACEAE 

Small epiphytic or terrestrial ferns mostly found in rainforests under very 
moist conditions. Rhizome long-creeping or short and thick, often slender or 
filiform, protostelic, clothed with hairs, sometimes without roots. Stipes unwinged 
or frequently winged almost to the base, mostly filiform or slender, glabrous or 
hairy. Lamina simple, bifid, palmatifid, 1-pinnate to pinnately decompound, 
usually 1 cell thick except the veins, membranous, almost transparent, lacking 
stomata. Pinnae with plain or pitted walls, glabrous or clothed with simple to 
elaborately branched hairs. Ultimate segments small, mostly 1-veined. Veins free, 
except in a few cases. False veins (if present) composed of rows of thickened cells 
which are not part of the vascular system. Sori marginal, terminal on the pinnules 
or on short lateral lobes. Involucre (indusium) bivalvate, tubular or obconic, the 
lips often much dilated. Receptacle columnar, included to long-exserted, bristle-like 
in the Trichomanes-group. Sporangia spherical to compressed, very shortly stalked 
or sessile, clustered along the receptacle, basipetal or maturing at the same time, the 
annulus complete, oblique or almost transverse, without definite stomial cells, 
opening by an obliquely lateral slit. Spores 32 to 420 in each spore, tetrahedral 
to globose, trilete, without a perispore, thin-walled, scabrate, granulate or 
subgranulate to papillate. Gametophyte ribbon-like and branching, thallose or 
filamentous, monoecious or rarely dioecious, green, mostly with the lateral cell-walls 
pitted, bearing discontinuous marginal cushions (in the HymenophyHum-group) or 
discontinuous archegoniophores (in the Trichomanes- group); germination rapid, of 
the 3-celled type (in the Hymenophy Hum-group) or the 4-celled type (in the 
Trichomanes- group) or rarely of a more complex type (in several species of 
Hymenophyllum subgen. Mecoclium); rhizoids occurring on the margin of the 
prothallus, usually branching and with thick brown walls; gemmae highly specialized 
and abundant; antheridia each with a 1-celled stalk, no funnel-shaped basal cell 
and no definite cap-cell, large and complex in the Hymenophyllum-group but smaller 
and simpler in the Trichomanes-group-, archegonia typically leptosporangiate, with a 
long, straight neck in the Hymenophyllum-group, the neck shorter (usually 4-cells 
long) in the Trichomanes-group. 

E. B. Copeland studied the Hymenophyllaceae in great detail, including his 
earlier work in monographs of Trichomanes sens. lat. and Hymenophyllum sens, 
lat. which were published respectively in Phil. Journ. Sci. LI (1933) 119-280, and 
Phil. Journ. Sci. LXIV (1937) 1-177. Later he proposed a new system of classification 
of this family in Phil. Journ. Sci. LXVII (1938) 1-110, recognizing 33 genera, and 
basing his revision partly on the earlier work by Presl and van den Bosch. Prantl, 
in Die Hymenophyllaceen (1875), also outlined a scheme of classification for this 
family. The usual practice has been to place the 600 to 650 species into two large 
genera, namely Hymenophyllum and Trichomanes, although some botanists have 
also recognized Cardiomanes (a monotypic genus from New Zealand with strong 
affinities to the Hymenophyllum- group of genera) and Serpyllopsis (a South American 
genus). However, Microtrichomanes, a group of about 10 species with reduced 
fronds, is the problem genus in the Hymenophyllaceae, as it stands half-way between 
the old comprehensive genera Hymenophyllum and Trichomanes. Although the 
involucre of Microtrichomanes is obconic (or rarely campanulate) as in the 
Trichomanes-group of genera, the teeth and bristle-like hairs of some species as 
well as the shape of the involucre (excluding the lips) suggest Sphaerocionium, 
especially S. lyallii (Hook, f.) Copel., as indicated by Copeland. 

It is clear to anyone studying the Hymenophyllaceae that the genera of the 
Trichomanes- group in Copeland’s classification are more sharply defined than those 
of the Hymenophyllum-group. Copeland himself remarked on this fact in Phil. 


2 


201 . Hymenophyllaceae. (Tindale) 


Journ. Sci. LXIV (1937) 5, stating that he had much greater trouble in the demarcation 
of natural groups in Hymenophyllum sens. lat. Of the Hymenophy llum-group of 
genera occurring in south-eastern Australia, Apleropteris and Sphaerocionium are 
sufficiently distinct but Mecodiwn , Meringium and Craspedophyllum are, in my 
opinion, linked with Hymenophyllum sensu Copeland in Genera Filicum (1948), 
so that I have treated them as subgenera of Hymenophyllum, as Copeland did in 
an earlier paper in Phil. Journ. Sci. LXIV (1937). Hymenophyllum is joined to 
Meringium by several dwarf species, viz. H. pumilum C. Moore (New South Wales), 
H. pumilio Ros. (New Caledonia) and H. minimum Rich. (New Zealand). It is 
difficult to ascertain by the degree of dissection of the involucre in which subgenus 
the 3 latter species with very reduced fronds should be placed. Also there is a 
small group of species in subgenus Meringium in which the laminal segments are 
almost entire, although they are serrated in most of the 60 species. Similarly, I 
have treated Mecodium as a subgenus of Hymenophyllum, since the line between 
them is difficult to draw. In this subgenus the segments of the lamina are entire 
except for 3 aberrant species (H. thuidium Harrington, H. reinwardtii van den Bosch 
and H. samoense Bak.), the last-mentioned species being found in tropical Queensland. 
Craspedophyllum has more claim to generic rank but the position is complicated 
by the possible relationship of the 2 Australian and New Zealand species to 2 species 
from South-eastern Asia. Hymenophyllum subgenus Meringium has some features 
which occur in the Trichomanes-group of genera, namely the elongated receptacles 
and the involucres which are partly valvate but tubular at the base. The cell-walls 
of the lamina are characterized by coarse-bordered pits in many species of subgenus 
Meringium, these being very similar to those of Selenodesmium. 

Copeland considered such features as a toothed laminal margin, stellate 
pubescence and supplementary wings on the rhachis as being advanced in the 
Hymenophyllum- group. He believed that the following were primitive characters 
in the Trichomanes-group: —filiform rhizome, absence of false veins and absence of 
any specialized thickening on the cell-walls of the lamina. He pointed out that 
all members of the Hymenophyllum-group are characterized by the first two features. 

Macroscopic features which are important in differentiating genera and 
species of the Hymenophyllaceae are as follows: — erect or creeping rhizome, winged 
or unwinged stipe and main rhachis, the type of hairs (stellate, forked or simple), 
the entire, crisped or toothed margin of the lamina, the shape and margin of the 
involucre, the included or long-exserted receptacle and the presence or absence of 
false and marginal veins. 

An important microscopic character of the Hymenophyllaceae is the thickening 
and pitting of the cell-walls. Elaborate drawings of these features may be seen in 
van den Bosch’s Hymen. Javan. (1861) and in Copeland’s monographs on Trichomanes 
and Hymenophyllum. Some are figured in this paper, e.g. No. 201, PI. IV and VI. 

The spores of the Hymenophyllaceae are trilete and tetrahedral, subtriangular 
or sometimes spherical, the spore coat being mostly thin and scabrate, subgranulate 
to granulate or papillate. The spores and young gametophytes within the spore 
coat are green due to numerous chloroplasts, being reminiscent of the spores of the 
Grammitidaceae. The Osmundaceae are also characterized by green spores but 
they are globose and have a smooth proximal region. The spores of the New 
Zealand species of the Hymenophyllaceae were treated in great detail by Harris in 
N.Z. D.S.I.R. Bull. 116 (1955) 73, plates 3 and 4. He observed that the spores of 
the Trichomanes-group in New Zealand with 2 exceptions were small (l.c. 73). On 
the whole the spores of the latter group are not as large as those of the Hymenophyllum- 
group in the Australian species. 


201 . Hymenophyllaceae. ( Tindale ) 


3 


As has been pointed out by a number of authors, the vegetative features of 
the Hymenophyllaceae are highly developed but the sori and sporangia are 
comparatively primitive as in the Gleicheniaceae. Although the sori are marginal 
in the Hymenophyllaceae instead of superficial as in the Gleicheniaceae, the sporangia 
of the filmy ferns have a complete, oblique or almost transverse annulus but without 
a definite stomium as in the higher ferns. 

Most of the Hymenophyllaceae are found in the rainforests of the tropics 
but others occur in the subtropical and moist temperate zones especially of the 
southern hemisphere, although only 2 species extend to Great Britain. Some species 
extend to southern Chile, the South Island of New Zealand and to Tasmania. Many 
of the Hymenophyllaceae are widely distributed especially Vandenboschia which 
Copeland has suggested, in Genera Filicum (1947) 37, is the most primitive genus 
of the Trichomanes-gtoup. There are 2 species of Vandenboschia in tropical 
Queensland but none occur in south-eastern Australia. 

The Hymenophyllaceae are well represented in Australia by 44 species of 
which 32 occur in the dense tropical rainforests of north-eastern Queensland. 
However, the filmy ferns are relatively common in the subtropical and temperate 
rainforests of south-eastern Australia where 18 species occur in moist shady situations, 
either as epiphytes on the trunks of trees and tree-ferns and on old logs, or (especially 
Hymenophyllum cupressiforme Labill.) growing on wet rocks in matted patches with 
mosses and hepatics. A few species, e.g. Hymenophyllum bivalve (Forst. f.) Sw. 
and Selenodesmium spp., sometimes occur as terrestrial plants amongst mosses on 
the rainforest floor especially near creeks. 

The lamina (other than the veins) of the Hymenophyllaceae is rarely more 
than one layer of cells in thickness, exceptions being Hymenophyllum australe Willd., 
H. dilalatum (Forst. f.) Sw. and H. demissum (Forst. f.) Sw. which are natives of 
Australia and/or New Zealand. The thin delicate fronds soon curl up in dry periods 
or if they are exposed to strong sunlight for a prolonged length of time. The 
shrivelled fronds of Hymenophyllum cupressiforme , which is probably the commonest 
filmy fern in New South Wales, are often seen on rocks at the edge of rainforests. 
This species is able to withstand short periods of drought and the laminas uncurl 
during the next rainy spell. The very thin, translucent lamina of the members of 
this family is unusual amongst the vascular plants. Unlike the epiphytes belonging 
to the Polypodiaceae and Davalliaccae, the Hymenophyllaceae appear to have no 
special methods of protection nor means of water storage in their rhizomes, roots 
or leaves. However, many filmy ferns (e.g. Hymenophyllum marginatum Hook, 
et Grev., of south-eastern Australia) occur in the mist regions on wet rocks or near 
waterfalls, so that the fronds are subjected to almost continually moist conditions. 

There is considerable overlap in the species occurring in south-eastern 
Australia (especially south of Gosford, N.S.W.) and in New Zealand. The following 
species are found in both regions: — Hymenophyllum flabellatum Labill., H. australe 
Willd., H. bivalve (Forst. f.) Sw., Polyphlebium venosum (R. Br.) Copel., 
Hymenophyllum peltatum (Poir.) Desv., H. rarum R. Br., Sphaerocionium lyallii 
(Hook, f.) Copel., Apteropteris malingii (Hook.) Copel. and Selenodesmium elongatum 
(A- Cunn.) Copel. The first 4 species extend to the subtropical rainforests of the 
North Coast of New South Wales and to Lamington National Park, south-eastern 
Queensland, whereas H. peltatum extends to the Bellenger River, N.S.W. S. lyallii 
and //. rarum have not been recorded north of the Blue Mountains, N.S.W., whereas 
A. malingii is confined to Western Tasmania but S. elongatum ranges from the North 
Coast of New South Wales to north-eastern Queensland. However, other species 
with a more tropical distribution do extend to south-eastern Queensland and the 


4 


201 . Hymenophyllaceae. ( Tindale ) 


North Coast of New South Wales, e.g. Microtrichornanes vitiense (Bak.) Copel., 
Cephalomanes atrovirens Presl, Gonoconmts saxifragoides (Pr.) van den Bosch, 
Macrog/ena caudata (Brack.) Copel. and probably *Microgonium tahitense (Nadeaud) 
Tindale. These species extend to either the Pacific Islands and/or Asia but do not 
occur in New Zealand. Of the filmy ferns found in north-eastern Queensland a 
a few are endemic e.g. Crepidomanes majorae (Watts) N. A. Wakef., f Crepidomanes 
walleri (Watts) Tindale, Hymenophyllum gracilescens Domin and H. kerianum Watts 
but others extend to Malaysia and the Pacific Islands. 

Cytologists have made chromosome counts of only a comparatively small 
number of the 600 to 650 species relegated to the Hymenophyllaceae. Certainly 
further counts would throw considerable light on the validity of Copeland’s genera. 
The cytological work on this family, cited in the text, has been mainly accomplished 
by Manton & Sledge, Mehra & Gurdip Singh, and Brownlie. There is a certain 
amount of overlap between members of this family occurring in New Zealand and 
in Eastern Australia, so that Brownlie’s counts for these species have been cited 
in the appropriate places. A few new counts for Australian species by the author 
have been added. 


1. Involucre distinctly bivalvate. Receptacle included or shortly exserted beyond the 
lips of the involucre in all the species from south-eastern Australia. 

2. Stellate or forked hairs (also some simple bristle-like hairs) present on the lamina 
or its margin. 

3. Typical lamina replaced by parenchyma prolonged into long papillae 

. Apteropteris I. 

3. * Typical lamina present . Sphaerocionium 2. 

2* Stellate or forked hairs absent on the lamina and its margin. 

4. Involucre without a definite tube, divided to or nearly to the base. Margin 
of the lamina entire or toothed. 

5. Margin of the lamina entire in the species from south-eastern Australia. 

6. Lamina with a black margin, minute, simple or once to thrice 

forked. Old cell-walls wavy-thickened ... 

. Hymenophyllum subgen. Craspedophyllum 4B. 

6.* Lamina without a black margin, pinnately divided. Cell-walls 

typically thin . Hymenophyllum subgen. Mecodium 4C. 

5.* Margin of the lamina toothed. Cell-walls straight or slightly wavy 

. Hymenophyllum subgen. Hymenophyllum 4A. 

4.* Involucre with a definite tube, bivalvate in the upper portion. Margin of 
the lamina toothed in the species from south-eastern Australia. Cell-walls 
usually thick and coarsely pitted . Hymenophyllum subgen. Meringium 4D. 

1.* Involucre tubular or obconic, the mouth entire or slightly 2-lipped, truncate or dilated. 
Receptacle long-exserted (except in Microtrichornanes and Crepidomanes walleri). 


* Footnote: Microgonium tahitense ( Nadeaud) Tindale comb. nov. (Basionym, Trichomanes 
tahitense Nadeaud, Enum. des Plantes Indigenes de Pile de Tahiti (1877) 18.) 

f Footnote: Crepidomanes walleri {Watts) Tindale comb. nov. (Basionym, Trichomanes walleri 
Watts in Proc. Linn. Soc. XXXIX, 1914 (1915) pi. lxxxvi, f. 3 a-d). 










201 . Hymenophyllaceae. (Tindale) 


5 


7. Rhizome long-creeping, 1.5 mm. or less in diam. Fronds mostly remote on the 
rhizome. Lamina soft. 

8. Stellate or forked hairs present on the margin of the lamina and its veins in 

the only species from south-eastern Australia. Sphaerocionium 2. 

8.* Stellate or forked hairs absent on the margin of the lamina and its veins in 


the species from south-eastern Australia. 

9. False veins present in the lamina. 

10. Fronds simple or lobed . Microgonium 5. 

10. * Fronds pinnate. Crepidomanes 6. 

9.* False veins absent in the lamina. 

11. Fronds palmate, flabellate to orbicular with a deeply incised 

margin. Axes sometimes proliferous . Gonocormus 7. 


11.* Fronds never palmate, flabellate nor orbicular with a deeply incised 
margin in the species from south-eastern Australia. Axes never 
proliferous. 

12. Fronds simple or dichotomous. Segments of the lamina 

1-veined or in simple fronds with the costa only. 

. Microtrichomanes 3. 

12.* Fronds pinnatifid to decompound. Segments of the lamina 
1-veined or pinnately veined. 

13. Segments of the lamina with pinnate veins .... Polyphlebium 8. 

13.* Segments of the lamina 1-veined. Macroglena 9. 

7.* Rhizome erect or oblique, tufted or very shortly creeping, stout, 1 cm. or more 

in diam. Fronds clustered on the rhizome. Lamina harsh or firm in texture. 

14. Fronds 1-pinnate. Cell-walls of the lamina relatively thin and wavy 

. Cephalometries 10. 

14.* Fronds decompound. Cell-walls of the lamina thick and coarsely pitted, 

qj* W avy . Sclctiodcstniutn 11. 

I. THE GROUP OF SPHAEROCIONIUM 

In this group 3 genera are considered to form a closely allied group, viz. 
Apleropteris, Sphaerocionium and Microtrichomanes. The fronds of Apteropteris 
have a stellate pubescence similar to those of Sphaerocionium but the anatomy of 
the lamina is more elaborate. Microtrichomanes with its shallow sori is obviously 
closely related to Sphaerocionium , since the gap between Sphaerocionium lyallii 
(Hook, f.) Copel. and Microtrichomanes palmatifidum (K. Mull.) Copel. is very 
narrow. The usually obconic involucre of S. lyallii is very similar to several species 
of Microtrichomanes (see page 7). The only species of Microtrichomanes occurring 
in south-eastern Australia, viz. M. vitiense (Bak.) Copel., is less morphologically 
similar to Sphaerocionium than most species of the former genus. Leptocionium, 
a monotypic genus from southern Chile, would also belong in this group. 

1. APTEROPTERIS Copel. 

Apteropteris Copel. in Phil. Journ. Sci. LXVIl (1938) 34; Copel., Gen. Fil. 
(1947) 34. 

HymenophyUum Sm. subgen. Apteropteris Copel. in Phil. Journ. Sci. LXIV (1937) 176. 

This is a monotypic genus found in Tasmania and New Zealand. Type of 
Genus: Apteropteris malingii (Hook.) Copel., (Basionym: Trichomanes malingii 
Hook.). 











6 


201 . Hymenophyllaceae. ( Tindale ) 


Apteropteris malingii (Hook.) Copel. in Phil. Journ. Sci. LXVII (1938) 35, 
pi. 1; Wakefield, Ferns Viet. & Tas. (1955) 4, fig. 9, 5. Habitat epiphytic and forming 
matted patches on the trunks and branches of Banksia and Arthrotaxis in temperate 
rainforests. Rhizome long-creeping, branched, smooth, brown, glossy, non-glaucous, 
cylindrical, filiform, wiry, pilose with stalked, stellate, ferruginous hairs. Stipes 
unwinged, filiform, 4 to 40 mm. long, distant on the rhizome, cylindrical, wiry, 
brown, smooth, clothed near the base with a tuft of stalked, stellate, ferruginous 
hairs and sparsely clothed on the middle and upper portions with similar hairs. 
Fronds including the stipes 2 to 20 cm. long. Lamina decompound, 1-2 to 8 cm. 
long and 0-7 to 2 cm. broad, very absorbent, grey or rusty-brown due to a dense 
tomentum of stellate trichomes each with 5 to 7 arms; instead of an ordinary lamina 
the parenchyma cells are prolonged into long narrow papillae containing chlorophyll 
and separated by air-spaces, the bases of the stellate hairs extending from the 
sclerenchyma up between the papillae. Primary pinnae 0-5 to 1-5 cm. long, 0-5 to 
1 cm. broad, the lower pairs ovate-rhomboidal, the upper more or less flabellate. 
Secondary segments cuneate or flabellate, deeply divided. Ultimate segments linear 
(12: 1 or more), terete, the apex obtuse. Sori indusiate, terminal on the segments. 
Involucre about 0-7 to 1 mm. long, orbicular-cupuliform, densely clothed with 
matted, stellate hairs, 2-valved to about two-thirds of the length, the valves denticulate 
at the apex. Receptacle included or slightly extruded, cylindrical. Sporangia 
compressed, slightly turbinate, sessile or almost so, the annulus composed of 14 to 
18 indurated cells and about 8 thin-walled cells. Spores globose-tetrahedral to 
globose, trilete, subgranulate, 37 to 51 jx x 37 to 51 |x. 

Distribution: Australia (Tasmania) and New Zealand (North and South Islands). 

Lectotype: On the mountains between Blind Bay and Massacre Bay, New Zealand, 
Christopher Maling (K, examined). Isotype? Golden Bay Ranges, Nelson, New Zealand, 
C. Maling (K, examined); Hooker considered this was possibly the same locality as that in which 
the lectotype was collected. This specimen consists of a single frond. Syntype: On the 
mountain range between Blind and Massacre Bay, Brunner (K, examined). I have chosen as 
lectotype the specimen which approximates most closely to the illustration and description by 
W. J. Hooker in Gard. Ferns (1862) pi. 64; also it was collected by Maling after whom the 
species was named. As in the plate, this specimen has a rhizome, whereas it is lacking in both 
of the other collections. 

Tasmania: West Coast: Mt. Roland, Sutton 1.1911 (NSW P1236); Cradle Mountain, 
Sutton 1.1911 (NSW P4966); ditto. Rodway 12.1915 (NSW P8277); ditto, Perrin 8.1932 (NSW 
PI237); ditto, 3,500 ft. alt., abundant on trunks of Arthrotaxis selaginoides in forest of the latter, 
Johnson 1.1949 (NSW P5615); Waldheim Forest, Cradle Valley, Ellis 5.1962 (NSW P8390); 
Macquarie Harbour, on Arthrotaxis selaginoides , Rodway 4.1906 (NSW P1238). 

Absolute Synonyms: Trichomanes malingii Hook., Garden Ferns (1862) pi. 64. 
Hymenophyltum malingii (Hook.) Mett., Hymen. (1864) 423, pi. 1, f. 32; Rodway, Tasm. Fl. 
(1903) 290; Copeland in Phil. Journ. Sci. LXIV (1937) 176. 

Apteropteris malingii is found in Western Tasmania at Cradle Mountain and near 
Queenstown in areas of very high rainfall. It is an epiphyte on the bark of Arthrotaxis 
selaginoides Don, A. cupressoides Don and Banksia sp. According to Crookes in New Zealand 
Ferns (1952?) 93, this rare mountain fern is chiefly found in New Zealand on the dead or dying 
trunks of Librocedrus bidwillii. 

As explained by Holloway in Trans. & Proc. N.Z. Inst. LIV (1923) 597, the structure of 
the lamina in Apteropteris is unique, as the narrow papillae containing chlorophyll take the 
place of the ordinary mesophyll. The lamina absorbs water from the atmosphere like a sponge 
but evaporation is very slow. 

Copeland suggested that Apteropteris is a derivative of Sphaerocionium, as indicated by 
the presence of stellate hairs on the fronds of both genera, although the anatomy of the lamina 
is very different. 


201 . Hymenophyllaceae. (Tindale) 

2. SPHAEROCIONIUM Presl 


7 


Sphaerocionium Presl, Hymen. (1843) 125; Copel. in Phil. Journ. Sci. LXVII 
(1938) 28; Copel., Gen. Fil. (1947) 34. 

Dermatophlebium Presl, Epim. Bot. (1849-52) 258; Copel. in Phil. Journ. Sci. LXVII 


HymenophyUum Sm. subgen. Sphaerocionium (Presl) Copel. in Phil. Journ. Sci. LXIV 
(1937) 164. 

Small epiphytic ferns with long-creeping, filamentous rhizomes. Stipes 
filiform, sometimes winged especially near the apex. Fronds flabellately or pinnately 
compound, clothed with branched, basally forked, stellate, bistellate or simple 
hairs on the axes, involucres and the margins of the lamina as well as on the surface 
of the lamina in other species; chromatophores minute and numerous; cell-walls 
thin. Veins free, with accessory wings not in the plane of the frond in some non- 
Australian species. Involucres rounded, 2-valved, usually cleft almost to the base, 
rarely obconic with an immersed tube (i.e. in S. lyallii). Receptacle shortly columnar, 
not projecting beyond the involucre. Sporangia sessile to very shortly stalked, 
more or less turbinate, the annulus broad, oblique to almost transverse. Spores 
globose to tetrahedral-globose, trilete, greenish. 

Type of Genus: Sphaerocionium hirsutum (Sw.) Presl (Basionym: 
HymenophyUum hirsutum Sw.), of Cuba, Mexico to Chile. See Copeland, Genera 
Filicum (1947) 34. 

This is a genus of about 63 species, although only 11 of them are found in 
the Old World. The majority of Sphaerocionium spp. (about 51 species) are confined 
to the rainforests of tropical America where they usually grow as epiphytes in the 
higher mountains. Only one species, viz. S. lyallii (Hook, f.) Copel., has been 
recorded from Australia, where it occurs on the Central Coast and Tablelands of 
New South Wales. 

So far the only chromosome count in Sphaerocionium is on one of the New 
Zealand species, viz. S. ferrugineum (Colla) Copel., for which Brownlie recorded 
n = 36 in Trans. Roy. Soc. N.Z. LXXXV (1958) 213. 

As indicated by Copeland in Phil. Journ. Sci. LXVII (1938) 29, the simple 
hairs of Sphaerocionium are always bristle-like and non-septate above the base or 
basal cell, as opposed to those species placed in HymenophyUum subgen. Meringium 
in which the simple hairs each consist of a row of cells. Microtriehomanes, a small 
genus to which Sphaerocionium is closely related, includes species with simple setae 
as well as some glabrous species which are unlike typical members of the genus 
Sphaerocionium (e.g. Microtriehomanes francii (Christ) Copel.). In Phil. Journ. Sci. 
LXVII (1938) 36, Copeland pointed out that the gap between S. lyallii and 
Microtriehomanes palmatifidum (K. Mull.) Copel. is very narrow. Amongst other 
features in common, both species are characterized by dark cells at the bases of the 
simple or branched hairs occurring on the laminal margin. The obconic involucres 
of M.parvulum (Poir.) Copel. are very similar to those of S. lyallii in shape and margin 
but hairs are absent. The fiabellate form of the lamina in S. lyallii resembles that 
of several species of Microtriehomanes including Af. palmatifidum. However 
Sphaerocionium is essentially characterized by branched hairs usually with a stalk 
cell, although a few simple hairs may also occur on the fronds. Future evidence 
may indicate that these 2 genera should be united under Sphaerocionium. 


8 


201 . Hymenophyllaceae. ( Tindale ) 


Morton in his critical revision of “ The American Species of Hymenophyllum 
section Sphaerocionium ” in Contrib. U.S. Nat. Herb. XXIX (1947) 142, divided 
this group into 2 subsections viz. subsect, ciliata and subsect, lanata. In the former 
subsection the hairs are borne only on the margins of the lamina and the veins, 
whereas in the latter they are also found on the surface of the lamina. The Australian 
and New Zealand species S. lyallii would be placed in subsect, ciliata according to 
this scheme. 


Sphaerocionium lyallii (Hook. /.) Copel. in Phil. Journ. Sci. LXVII (1938) 
33; Crookes & Dobbie, New Zealand Ferns, ed. 5 (1952?) 89, 90 (photograph). 

Habitat epiphytic or rupestral, forming matted patches on trees and rocks 
in subtropical rainforests, usually between 1,200 and 3,500 ft. altitude in New South 
Wales. Rhizome long-creeping, branched, black or very dark brown, slightly 
glossy, filiform, about 0T to 0-2 mm. in diam., sparsely pilose with light red-brown, 
simple hairs (composed of 2 to 4 cells) or clothed with once forked hairs. Stipes 
not winged, 0*1 to 2-5 cm. long, filiform, about 0 1 mm. or less broad, distantly 
spaced on the rhizome, black or very dark brown, slightly glossy, clothed at the 
base with a tuft of long, pale ferruginous hairs and sparsely pilose towards the middle 
and apex with easily abraded, light ferruginous, simple or once or twice forked 
hairs. Fronds not dimorphic. Lamina divided flabellately or digitately almost to 
the base of the frond into dichotomous segments, very rarely simple, very 
membranous, pellucid, pale green when fresh, brown when dried, 3 to 16 mm. long, 
3 to 16 mm. broad, deltoid or broadly obovate (6: 5) to very broadly obovate (6: 6), 
rounded (6: 5) to almost orbicular (6: 6) or sometimes once forked in small fronds, 
the internal cell-walls slightly wavy-thickened. Ultimate segments 1 to 16, sessile, 
opposite, plane, ascending, oblong (2: 1) to cultrate (10: 1), 1-5 to 10 mm. long, 

1 to 1 -5 mm. broad, the surface glabrous, the apex rounded, truncate or emarginate, 
the margin bearing small persistent teeth with fugacious, reddish, simple or more 
usually once or twice forked hairs (the apical cell of each branch bent) or occasionally 
with basally forked hairs. Veins free, dichotomous, black or very dark brown, 
glabrous or with a few, forked, reddish hairs. Sori indusiate, marginal, solitary 
or very rarely in pairs at the apex of the ultimate segments, deeply sunken in the 
lamina. Involucre 1 to 1-5 mm. long, 0-8 to 2-5 mm. broad, valvate, obcordate or 
cuneate-obconical, green; sometimes 2-lipped, the mouth not dilated nor with a 
specialized border; apex retuse, erose or toothed, each tooth bearing a reddish, 
forked or basally twice forked hair. Receptacle included, cylindrical. Sporangia 
few, sessile, compressed, slightly turbinate; annulus very broad, almost transverse, 
composed of 16 to 19 indurated cells and ca. 9 thin-walled cells, dehiscing 
obliquely laterally. Spores subtetrahedral to globose, trilete, finely granulate, green, 
equat. diam. 24-38 to 39-75 g x polar diam. 22-5 to 37-5 g. (No. 201, PI. I.) 

Distribution: New Zealand and Australia (New South Wales, mainly in the Blue 
Mountains and the ranges of the lower Central Coast). 

Holotype: Thomson’s Sound, South West Coast, Middle Island, New Zealand, on trees, 
D. Lyall 3.1851 (K), examined. 

New South Wales: Blue Mountains, Betche 8.1895 (NSW P593); Kings Tableland, 
Forsyth 2.1903 (NSW P2212); Wentworth Falls, in deep gullies, Betche 8.1895 (NSW PI860); 
ditto, Forsyth, probably about 1900 (NSW P2110); ditto. Chapman 12.1909 (SYD); Katoomba, 
Forsyth 12.1899 (NSW P2113); Bulli, Kirton (NSW P2207); Minnamurra Falls, about 3 miles 
W of Jamberoo, 1,200 ft. alt., in rainforest, in deep ravine above creek on boulder, in the mist 
region, Judd 5.1954 (NSW P6791); above Minnamurra Falls, 2,200 ft. alt., on old trees of 
Ceratopetalum apetalum, Judd 5.1955 (NSW P7161); pass above Kiama, Moore 2.1890 (NSW 
P2111, P2112); ditto, Harwood 4.1895 (NSW P2206); Brogher’s Creek, Kangaroo Valley, 
Bauerlen 8.1880 (NSW P2960); Mt. Cambewarra, Watts 10.1907 (NSW P2211, P1859). 


201 . Hymenophyllaceae. ( Tindale ) 


9 


Absolute Synonyms : Hymenophyllum lyallii Hook, f., FI. N.Z. II (1854) 16. Trichomartes 
lyallii (Hook, f.) Hook, in Hook. & Bak., Syn. Fil. (1867) 77; Copel. in Phil. Journ. Sci. LI 
(1933) 163, pi. 7, f. 7, pi. 11 f. 4. 

Equivalent Synonyms: Trichomanes calvescens van den Bosch, Ned. Kruid. Arch. V 
(1863) 199; Hook. & Bak., Syn. Fil. (1868) 77, Holotype by Monotypy: New South Wales, 
Vicary 1836-7 (K), examined. This specimen was labelled T. calvescens n.sp. by van den Bosch. 
Trichomanes digitatum Sw. var. calvescens (v.d. Bosch) Domin in Bibl. Bot. LXXXV 
(1913) 13; Maiden & Betche, Census N.S.W. PI. (1916) 1; Melvaine in Proc. Linn. Soc. N.S.W. 
LXI (1936) 113. I do not consider that Bentham implied this combination in FI. Austr. VII 
(1878) 702, as was suggested by Maiden and Betche, although this transfer was made by Domin 
in Bibl. Bot. LXXXV (1913) 13. Holotype: the same as for T. calvescens. 

This species was misidentified as Trichomanes digitatum auctt. non Sw. in Benth., FI. 
Austr. VII (1878) 702 and Moore & Betche, FI. N.S.W. (1893) 503. For many years in New 
South Wales Microtrichomanes digitatum (Sw.) Copel. (syn. Trichomanes digitatum Sw.) was 
confused with Sphaerocionium lyallii but M. digitatum may be distinguished from the latter by 
the unbranched, bristle-like hairs on the margins of the lamina and by the simple, ferruginous 
hairs on the rhizomes. Hymenophyllum cattlettii (cattletti ) C. Moore et Betche, Handb. FI. 
N.S.W. (1893) 503, nom. illegit., was published in synonymy under T. digitatum but I have 
examined a specimen so-labelled by Moore at the Kew Herbarium. This material, which was 
collected in the lllawarra by C. Moore and received at Kew in September 1890, agrees well with 
Australian specimens of S. lyallii. 

On the whole the fronds of the New Zealand specimens of S. lyallii are more robust, the 
involucres slightly larger and more retuse, also the hairs on the lamina and involucres are usually 
more complex. In the material from New Zealand twice forked and stellate hairs are much 
more common on the fronds than in the New South Wales specimens. The number of ultimate 
segments of the fronds is usually greater in the New Zealand material, i.e. 18 to 26 instead of 
1 to 16. However, a specimen from the D.S.I.R., Wellington, New Zealand, collected at the 
Waitakere Ranges, North Auckland, by Carse No. 2165 in January, 1919, is very similar to 
Australian material of this species. It has from 10 to 20 ultimate segments on each frond. The 
involucres are folded back but are usually straight or almost so, instead of being retuse, also 
simple hairs are found on the apices of the involucres. Material collected on the eastern side 
of the Waitakere Range by Crookes in March 1962 (NSW P8441) approximates closely to the 
Australian material but NSW P8440 obtained in the same locality by Crookes & Tindale in 
January, 1962, although characterized by few laminal segments, i.e. 5 to 14, has very large sori 
which are up to 3 mm. broad. 

The hairs on the margins of the laminas in the holotype of S. lyallii are forked, twice 
forked, basally forked, stellate or rarely simple. The retuse involucres have an apical row 
of mainly forked hairs, also there are 14 to 25 ultimate segments in the fronds. On the other 
hand the holotype of Trichomanes calvescens van den Bosch is typical of the Australian material, 
the hairs on the fronds and involucres being either simple (consisting of 2 to 4 cells in length) or 
rarely once forked. Some of the involucres on this specimen are retuse. 

When discussing the development of the gametophyte in New Zealand material of 
S. lyallii, Atkinson, in Phytomorphology X (1960) 32, mentioned that the “ second wall in the 
spore is suppressed as well as one or two divisions in the arms ” but this condition occurs 
infrequently. 


3. MICROTRICHOMANES (Mett.) Copel. 

Micro-trichomanes Mett., Hymen. (1864) 413, as a Group of Trichomanes. 
Microtrichomanes (Mett.) Prantl, Hymen. (1875) 51, as a Sect, of Gonocormus; 
Copel. in Phil. Journ. Sci. LI (1933) 153, as a Group of Trichomanes', C. Chr., 
Ind. Fil., Suppl. 3 (1934) 187, as a subgenus of Trichomanes', Copel. in Phil. Journ. 
Sci. LXVII (1938) 35, as a genus, also Copel., Gen. Fil. (1947) 34, and Fern FI. 
Phil. I (1958) 55. 

Small, fragile epiphytes. Rhizome filiform, creeping. Stipes widely spaced 
on the rhizome, not winged throughout. Axes winged. Lamina usually 
dichotomous, sometimes simple by reduction, mostly brownish, false veins absent, 
without veinlets except the costae of the segments, the margin not specialized but 


10 


201 . Hymenophyllaceae. ( Tindale ) 


often bearing cilia or setae, the cell-walls thin. Sori terminal on the longer segments 
of the lamina. Involucre usually obconic or rarely campanulate, entire, winged to 
the apex. Receptacle mostly shortly exserted, slender. Sporangia large, almost 
spherical. Spores tetrahedral to almost globose, trilete, green. Gametophyte 
3-celled llymenophyllum- type. 

Type of Genus: Microtrichomanes digitatum (Sw.) Copel., based on 
Trichomanes digitatum Sw., of Mauritius, Borneo, Java, the Philippines, Malaya 
and perhaps Samoa. 

According to Copeland this is a small genus of about 10 species occurring 
in the Southern Hemisphere. He asserted that Microtrichomanes was derived 
from Sphaerocionium by reduction of the lamina. The involucre of the latter genus 
is mostly valvate instead of obconic or rarely campanulate as in Microtrichomanes, 
also the laminal hairs in Sphaerocionium are almost always branched not simple. 
However, the brownish lamina, the hairs and teeth in some species and the shape of 
the involucre (excluding the lips) of Microtrichomanes indicate relationship with 
Sphaerocionium. The short lips of the involucre and the shortly extruded receptacle 
denote affinities with the Trichomanes- group of genera but Microtrichomanes is 
clearly intermediate between this group and Hymenophyllum sens. lat. There is 
only one species of Microtrichomanes in south-eastern Australia. 


Microtrichomanes vitiense ( Bak .) Copel. in Phil. Journ. Sci. LXVII (1938) 
37; Copel. in Journ. Arn. Arb. XXX (1949) 433. 

Habitat in tropical or subtropical rainforests forming densely matted patches 
on trees and logs. Rhizome elongated, filiform, long-creeping, branched, densely 
clothed with brown, linear, unbranched hairs each composed of a single row of 
cells. Stipes winged towards the apex or not winged, 1 to 2 mm. long, densely 
tomentose with similar hairs to those on the rhizome. Lamina simple or once 
forked, membranous, translucent, glistening, mostly yellowish-green, narrow oblong 
(3: 1) to oblong (2: 1) or somewhat elliptical (2: 1), 2 to 11 mm. long including 
the stipes, T5 to 4 mm. broad; margin entire or obscurely sinuate, somewhat 
crisped in dried specimens; apex emarginate, rounded or truncate; base tapering 
gradually or abruptly; one-veined unless the frond is forked, no lateral veins 
present; costa black and clothed especially near the apex with a few, appressed, 
oblong, rather bulbous, opaque or reddish hairs 1 to 3 cells long and with red joints 
between the cells; cell-walls of the lamina very thin and hyaline, the contents 
aggregated densely against all the lateral walls except those of the margin. Sori 
indusiate, terminal, 1 to 2, deeply sunken in the apex of the fronds or its lobes. 
Involucre somewhat broadly trumpet-shaped, urceolate-oblong, clothed especially 
towards the base with hairs of the same type as on the costa; rim scarcely dilated 
but folded in dried specimens or sometimes reflexed, entire or slightly fluted, 
obscurely 2-lipped, equal to the breadth of the lobe. Receptacle oblong, scarcely 
or not exserted, rarely long-extruded. Sporangia subsessilc, almost spherical, 
with an oblique annulus of 14 to 19 indurated cells and 2 or 3 thin-walled cells. 
Spores tetrahedral to globose, trilete, green, granulate, equal, diam. 33-75 to 37-5^- 
X polar diam. 30 to 33-75|x. (No. 201, PI. V, f. 1.) 

Distribution: Fiji, New Caledonia and Eastern Australia (Queensland to the North 
Coast of New South Wales). 

Holotype: Fiji, Milne (K), examined. 

Queensland: Herberton, Evelyn Scrub, R. F. Waller 9.1909 (NSW P2116); near Tully 
Falls, Watts 7-8.1913 (NSW P3783, P3767); Yandina, Simmonds 3.1891 (BRI); Maroochy 


201 . Hymenophyllaceae. ( Tindale) 


11 


River, Bailey 10.1874 (BRI); Brisbane, Bancroft 3.1894 (NSW P2118); Brisbane River, Bailey 
3.1873 (K.); Three Mile Scrub, Brisbane, on stems of trees, Bailey (BRI); Brookfield, near 
Brisbane, Bailey 12.1888 (BRI); Running Creek, Lamington Glen, epiphytic on Ficus 
stephanocarpa in rainforest on the edge of a creek, Goy & Smith No. 20, 10.1937 (BRI). 


New South Wales: Foot of Mt. Lindsay, Forsyth 9.1900 (NSW P2114); Richmond 
River, Mueller, received 9.1879 and 10.1881 (K); Richmond River, Watts 5.1899 (NSW P3766) 
and 1906 (NSW P3768); Binnaburra Road, Richmond River, on a tree, Watts 5.1899 (NSW 
P3762); North Creek Road, Watts 4.1899 (NSW P3763); Uralba, Richmond River, Watts 
1898 (NSW P3787)- Teven, Richmond River, Watts 4.1900 (NSW PI 141); Marshall’s Falls, 
Alstonville, Bauerlen 5.1891 (NSW PI857, PI858); Maxwell’s Scrub, Pearce’s Creek, Richmond 
River, Watts 9.1896 (NSW P3765). 


Absolute Synonym: Trichomanes viliense Bak. in Journ. Linn. Soc. Bot. IX (1866) 338, 
t. 8D; F. Muell., Fragm. Phyt. Austr. VIII (1873) 32; Benth., FI. Austr. VII (1878) 701; F. M. 
Bail., Lith. Ferns Queensl. (1892) t. 21; Moore & Betche, Handb. FI. N.S.W. (1893) 503; 
F. M. Bail., Queensl. FI. (1902) 1941; Domin in Bibl. Bot. LXXXV (1913) 10, t. 3, f. 3; Maiden 
& Betche, Census N.S.W. PI. (1916) 1; Copel., Ferns of Fiji, B.P. Bish. Mus. Bull. No. 59 (1929) 
24; Copel, in Phil. Journ. Sci. LI (1933) 157, pi. 9, f. 1-2; C. Chr., Ind. Fil., Suppl. 2 (1934) 
193; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114. 


The holotype of M. vitiense was collected in Fiji by Milne but there is insufficient difference 
between the Australian and Fijian material for them to be regarded as 2 species. The holotype 
is characterized by a relatively narrow involucral tube but so have several specimens from 
New South Wales, e.g. NSW P3765 (collected at Maxwell’s Scrub, Richmond River) which was 
matched with the type. No differences were noted in the internal or marginal cell-walls of the 
Fijian and the Australian material. The somewhat bulbous hairs which occur at the base of 
the involucre and also along the midrib of the lamina especially near the apex, are similar in 
the Fijian and Australian specimens. These hairs are usually 1 to 3 cells in length, mostly 
appressed, with rounded apices and opaque or more usually reddish with dark red walls. The 
holotype has dark green fronds, as is also the case in NSW P2118 which was collected at Brisbane 
by Bancroft in March 1894, but other specimens have very light yellowish-green fronds, e.g. 
NSW P318 which was collected at Viti Levu, Fiji by Degener No. 14607. The fronds are 
bilobed in the material from Viti Levu and also in some Australian specimens but several 
collections have simple fronds only. The fronds of Degener No. 14607 from Viti Levu are 
larger than those of most of the fronds in the holotype, also the rim of the involucre is more 
pronounced and expanded. However similar involucres are found in the following Australian 
specimens:—Richmond River, New South Wales ex Herb. F. Muell. (K) and Brisbane, Bancrolt 
3.1894 (NSW P2118). 


I consider that Trichomanes powellii Bak. should be kept as a distinct species, since the 
fronds range from simple to eight times forked, the “cuff” of the involucre is proportionally 
broader, also the involucre is usually separate from the apex of the frond and projects beyond it. 
The rim is incipiently 2-lipped and the receptacle included or shortly excluded as in Rechmger 
No. 3379, collected at Upolu in Samoa in August, 1905 (K). However Christensen in Bish. 
Mus. Bull. No. 177 (1943) 13, stated that the involucre in T. powellii is cyathiform-obcomc 
with a rather deeply 2-lipped mouth, the lips entire, round, large, spreading or subieflexed at 
maturity.” The fawn or reddish hairs on the midrib and involucral bases are alike tnV. vt tense 
and T. powellii , also the structure of the internal and marginal cells is similar. The rcflexed 
involucral lips in M. vitiense are narrower and the whole plant is much smaller l.e. 0- to I cm. 
long instead of 1 to 1-8 cm. long as in T. powellii. At tht kew Herbarium I examined the 
holotype of T. powellii which was collected in Samoa by the Rev. T. Powell and was received by 
Baker in May, 1866. 


Microtrichomanes vitiense is closely allied to Microgomum motley, van den Bosch which 
differs from the former in having lateral veinlets from the costa and deeply cordate sterile fronds 
also the greater part of the involucral tube projecting beyond die laming M was 

first recorded from north-eastern Queensland by Domin in Bibl. Bot. LXXXV (1)13) 9, but it 
is also found in the Philippines, Malaya, Ceylon Borneo, Java and New Guinea if this species 
is construed in the broad sense to include Trichomanes beccananum Cesati. M. mot ley i has 
false veinlets (in addition to true veinlets) in its laminas but they are absent in Microtrichomanes 
vitiense where the costa is the only vein. 


Amongst specimens sent to me recently by Dr. H. Hurlimann was a collection of M. 
vitiense from New Caledonia, obtained by Baumann in April, 1951, on the summit crest ot 
l’Oua Tilou 1 000 to 1,100 m. (Expedition botanique en Nouvelle Caledonie 1950-52, Mission 
Franco-Suis’se No. 12420). This appears to be the first record of M. vitiense from New Caledonia. 


12 


201 . Hymenophyllaceae. ( Tindale) 


II. THE GROUP OF HYMENOPHYLLUM 

In south-eastern Australia the HymenophyHum-group is considered to consist 
of a single large genus viz. Hymenophyllum with 4 subgenera represented in this 
region. The latter are as follows:— 

A. Subgenus Hymenophyllum (3 species in south-eastern Australia). 

B. Subgenus Craspedophyllum (1 species in this region). 

C. Subgenus Mecodium (3 species in this region). 

D. Subgenus Meringium (1 species in this region). 

The relationship of these subgenera is discussed on page 2. 


4A. HYMENOPHYLLUM Sm. SUBGENUS HYMENOPHYLLUM 

Hymenophyllum Sm. subgen. Euhymenophyllum sensu Copel. in Phil. Journ. 
Sci. LXIV (1937) 77. 

Copeland later considered this group of species to be of generic rank, i.e. Hymenophyllum 
sens, strict, in Phil. Journ. Sci. LXVII (1938) 37, and in Genera Filicum (1947) 34. 

Small, matted, epiphytic or terrestrial ferns. Rhizome creeping, narrow. 
Lamina pinnately decompound, the margins of the segments minutely denticulate 
or spinulose-dentate, the teeth falcate or straight, glabrous; cell-walls pitted, 
scarcely or not differentiated. Sori terminal on the ultimate segments or close to 
the main rhachis on short branchlets (1 to 3 in a row) on the upper side of each 
secondary rhachis. Involucre divided to the middle or to the base into 2 lips which 
are toothed or smooth. Receptacle included or very slightly exserted. Sporangia 
large, sessile. Spores tetrahedral to almost globose, trilete, papillate to granulate. 

Type of Subgenus: Hymenophyllum tunbridgense (L.) Sm. (Basionym: 
Trichomanes tunbridgense L.). 

In the restricted sense this is a subgenus of about 25 species principally 
occurring in the temperate parts of the world. Very few species extend to the tropics 
but a number occur in Japan, Norway, Scotland, New Zealand, Chile, South Africa 
and the Antarctic Islands. Four species are represented in Eastern Australia, the 
commonest being H. cupressiforme Labill. which occurs in subtropical and temperate 
rainforests from south-eastern Queensland to Tasmania. H. peltatum (Poir.) 
Desv. is found at comparatively high elevations in New South Wales and Victoria 
as well as in the mountains of Tasmania and along the rivers of the latter State near 
sea level in mixed Nothofagus forest. Hymenophyllum gracilescens Domin is found 
in the tropical, mountain rainforests of north-eastern Queensland. 

Chromosome counts have been made for a number of species of subgen. 
Hymenophyllum, the base numbers being 9, 11 and 13. Manton recorded counts 
of n = 13 for II. tunbridgense (L.) Sm. and n = 18 for H. wilsonii Hook, (as II. 
unilaterale Bory) in “ Problems of Cytology and Evolution in the Pteridophyta ” 
(1950) 272-4, fig. 274 a and b. Brownlie has made counts for 2 members of this 
group, viz. n = 11 for material of H. peltatum collected in New Zealand and n = 22 
for H. revolutum Col. (endemic to New Zealand). These were published in Trans. 


201 . Hymenophyllaceae. ( Tindale ) 


13 


Roy. Soc. N.Z. LXXXV (1958) 213. The author obtained a count of 2n = 22 
for Tasmanian material of H. peltatum and n = ca. 24 for specimens of 
H. cupressiforme Labill. collected in New South Wales (NSW P8425). 

There are 3 species of Hymenophyllum sens, strict, in south-eastern Australia. 


1. Sori terminal on the ultimate segments of the lamina, borne at the apex of the frond 

... H. pumilum 1. 

1.* Sori borne on short, specialized, lateral branches of the primary pinnae usually on 
the acroscopic side. 

2. Involucre entire, narrowly ovate to broadly ovate or broadly oblong, dark 
brown in the older stages, the apex obtuse or often pointed. Sori usually 2 or 3 
in a row on short lateral branches on the upper surface of the primary pinnae, 
sometimes solitary. Primary pinnae branching on the upper side only .. H. peltatum 2. 

2.* Involucre toothed, undulate or sometimes almost entire, suborbicular or 
depressed, transversely ovate with a short, cuneate tube, green in the older 
stages, the apex broadly rounded. Sori usually solitary on short, lateral branches 
on the upper surface of the primary pinnae, rarely up to 6 and extending to the 
lower surface. Primary pinnae branched on both sides . H. cupressiforme 3. 


1. Hymenophyllum pumilum C. Moore in Hook. & Bak., Syn. Fil. (1874) 
464; Benth., FI. Austr. VII (1878) 706; Moore & Betche, Handb. FI. N.S.W. 
(1893) 504; C. Chr., Ind. Fil. (1905) 366; Domin in Bibl. Bot. LXXXV (1913) 
25; Maiden & Betche, Census N.S.W. PI. (1916) 1; Melvaine in Proc. Linn. Soc. 
N.S.W. LXI (1936) 114; Copel. in Phil. Journ. Sci. LXIV (1937) 91 (excluding 
H. moorei Bak. which also comprises pi. 45 f. 1-3); l.c. LXVII (1938) 39, 44. 

Habitat epiphytic in subtropical rainforest in the mountains, occurring in 
densely matted patches. Rhizome long-creeping, branched, very dark brown or 
black, about 0T mm. in diam., sparsely clothed with a few, dark red-brown, simple 
hairs consisting of a single row of cylindrical cells. Stipes not winged except for 
an almost imperceptible wing towards the apex, 4 to 17 mm. long, filiform, distant, 
very dark brown or black, wiry, minutely scabrous, glabrous except for a few, basal 
hairs of the same type as on the rhizome. Rhachis broadly winged or rarely terete 
and not winged between the lowest pairs of primary pinnae, zig-zagged, black, 
glabrous or with an occasional, red-brown, linear or oblong, unbranched hair 
consisting of a single row of cells. Lamina fiabellately pinnate-pinnatifid, 
membranous, pellucid, brownish-green when dried, irregular in outline, very broadly 
ovate or somewhat rhomboidal, 0-7 to 1-8 cm. long and 0-5 to 1-5 cm. broad, not 
dimorphic, glabrous, the internal cell-walls thick and pitted. Primary pinnae 
fiabellately divided at the base into 2 to 4 segments, the upper usually simple. 
Ultimate segments mostly narrowly oblong (3: 1) to broadly oblong (3:2), 2 to 
9 mm. long, 1-3 to 2 mm. broad, the apex rounded or more rarely truncate or 
emarginate, the margin spinulose-dentate, the teeth distant, often dark brown or 
black, consisting of about 3 cells on a broad base. Veins black, forked once in 
each segment, not reaching the margin, clothed with an occasional, dark red, simple 
hair consisting of 3 to 7 cells. Sori indusiate, terminal on the ultimate segments 
at the apex of the fronds, 1 to 2 per frond, free, projecting beyond the segments, 
not immersed, up to twice as broad as the ultimate segments but always broader 
than the latter. Involucre 1*5 to 2 mm. long, 0-9 to 1-5 mm. broad, valvate, divided 
to the middle or almost to the base into 2 lips, rounded or oblong, greenish-brown, 
glabrous; apex broadly rounded with a minutely spinulose-dentate margin, the 
teeth small, obtuse and irregular in size, not occurring on the sides of the involucre; 
base cuneate; marginal row of cells elongated but scarcely specialized. Receptacle 
thickened towards the base, included. 




14 


201 . Hymenophyllaceae. ( Tindale ) 


Distribution: New South Wales, Australia (mainly the Blue Mountains and the pass 
above Kiama). 

4t Holotype: Mill Creek, Mt. Tomah, N.S.W., C. Moore, received 6.1871 (K), labelled 
“ Hymenophyllum pumilum Moore ” in his own handwriting. 

New South Wales: Mill Gully, Mt. Tomah, Carron (NSW P1907); Mt. Tomah 
(spelled Thomar), Blue Mountains (K), labelled by C. Moore; Sydney, Blue Mountains, 1837, 
ex Herb. T. Moore (K), (“ Sydney ” covered a wide area at that period); the pass above Kiama, 
C. Moore 2.1890 (NSW P7182). 


In Moore’s original description of H. pumilum the margin of the involucre is said to be 
entire but the valves of the holotype were found to be minutely spinulose-dentate on examination 
with a lens. 

According to Copeland in Phil. Journ. Sci. LXIV (1937) 91, Hymenophyllum moorei 
Bak. of Lord Howe Island, is a synonym of H. pumilum. However, after an examination of a 
number of specimens from Mt. Gower and Mt. Lidgbird, Lord Howe Island, as well as the 
types of both species at the Kew Herbarium in 1950, 1 consider that they are distinct species 
since the involucre of H. moorei is larger, i.e. 2 to 3 mm. long and 1-5 to 2 mm. broad, also the 
apex of the involucre is more deeply toothed, i.e. fimbriate or sometimes with hooked teeth. 
These teeth rarely occur on the sides of the involucre. The sori of H. moorei are rarely terminal 
but mostly lateral, stalked and sunken between 2 segments. The hairs of the rhizome and the 
laminal teeth of these 2 species are similar, although these teeth are longer in H. moorei. In 
Pacific Science XIV (1960) 244, Brownlie also maintains these 2 species as distinct after examining 
the holotypes of both taxa. 

Another closely allied, dwarf species is Hymenophyllum minimum A. Rich. (syn. Meringium 
minimum (A. Rich.) CopeL) which is found in New Zealand and the Auckland Islands. It has 
small, flabellately divided fronds and very large, terminal, stalked involucres with hooked teeth 
oi fimbriae along the margin and on the back of the valves. There are also spinulose teeth on 
the margin of the lamina. 


2. Hymenophyllum peltatum ( Poir .) Desv., Prodr. (1827) 333; Melvaine 
m Proc. Linn. Soc. N.S.W. LXI (1936) 114; Copel. in Phil. Journ. Sci LXIV (1937) 
Jin*?!' ^' c ‘ LXVII (1938) 38; Dobbie & Crookes, New Zealand Ferns, ed. 5 

(1952) 98, 99 (photo); N. A. Wakefield, Ferns of Victoria and Tasmania (1955) 6, 

4 i. 6. 


Habitat epiphytic on trees or fallen logs of Nothofagus cunninghamii and 
Atherosperma moschatum or growing on rock faces or sometimes on granite or 
dolerite boulders, from sea level to 5,300 ft. alt., sometimes in mixed Nothofagus 
forest along rivers. Rhizome filiform, wiry, long-creeping, much branched, ca. 
0'2 mm. in diam., clothed with brown, undivided hairs which are 2 to 4 cells long. 
Stipes filiform, wiry, widely spaced on the rhizome, 0-6 to 6 cm. long, ca. 0-2 mm. 
in diam., with an almost imperceptible wing towards the apex. Main rhachis with 
a narrow wing which broadens towards the apex, zig-zagged, glabrous. Fronds 
L5 to 17 cm. (usually 3 to 8 cm.) long including the stipe. Lamina 1-pinnate- 
pinnatifid to 2-pinnate-pinnatifid, membranous when young, herbaceous in the 
older stages, dark green, narrowly elliptical (4: 1) to elliptical (2: 1), lanceolate 
(3: 1), cultrate (6: 1) to linear (12: 1), 1-5 to 10 cm. long, 0-5 to 1*8 cm. broad; 
the laminal cells ornamented, the cell-walls pitted. Secondary rhachis with an 
occasional simple hair. Primary pinnae alternate towards the base and opposite 
near the apex or usually alternate throughout, unilateral, digitately pinnatifid. 
Ultimate segments 3 to 6 mm. long, 0-5 to 1 mm. broad, cultrate (6: 1) to narrowly 
oblong (3: 1), developed on the upper side of the primary pinnae, the apex obtuse 
or truncate, the margin spinulose-dentate. Veins free, forked but undivided in 
each ultimate segment, not reaching the margin. Sori numerous, usually occurring 
towards the apex of each frond, 1 to 3 borne on the upper basiscopic side of the 


201 . Hymenophyllaceae. ( Tindale ) 15 

secondary rhachises, each borne on a stalk up to 1 mm. long. Involucre narrowly 
ovate to broadly ovate or broadly oblong, 0-5 to 1 mm. broad, 1 to 1-8 mm. long, 
often pointed, dark brown and woody in the old condition but green in the early 
stages, divided to the middle or below the middle into 2 entire valves, cuneate at 
the base. Receptacle included. Sporangia subsessile, slightly turbinate, with an 
oblique annulus composed of 18 to 24 indurated cells and about 6 thin-walled cells. 
Spores subtriangular, trilete, with long narrow laesurae, finely papillate equat. 
diam. 43-13 to 50-63|x X polar diam. 37-5 to 41 -25[x. 

Distribution: Mascarene Islands, Tristan da Cunha?, New Zealand, Australia (New 
South Wales, Victoria and Tasmania) and South Africa. 


Holotype: lie de France (Mauritius), Bory de Saint-Vincent (P), examined. 

New South Wales: Head of Bellenger River, 4,500 ft. alt., on trees and rocks Herb 
Mueller (MEL); Barrington Tops, Coast Range, Boorman 1.1916 (NSW P1234, P1905 P2906)’ 
Thredbo River, Mt. Kosciusko, Maiden & Forsyth 1.1899 (NSW P1232); Thredbo River Gorge" 
Mt. Kosciusko, on rock face in gully, not common, 5,300 ft. alt., Johnson & Constable 1.1951 


Victoria: Wotan Creek, $ mile below Cleve Cole Hut, southern face of Mt Bogong 
(ca. 5,800 ft.), on sheltered damp rock faces, not uncommon, Willis 1.1947 (MEL); Mt. Bostons 
6,500 ft. alt., Tadgell 2.1923 (MEL); Bogong and Fainter High Plains, 5,500-6,000 ft alt 
Tadgell 2.1923 (MEL); head of Catherine River, Mt. Speculation, Barry Mts., 5,300 ft. alt ’ 
in rocky gulches, uncommon, Willis 1.1945 (MEL); head of Tyers River, near Mt. Mueller’ 
Baw Baw Plateau, amongst granite boulders at ca. 4,500 ft. alt., Willis 3.1951 (MEL); head of 
Taggerty River, Lake Mountain, Singleton 9.1943 (MEL); head of Taggerty River near 
Marysville, at about 4,000 ft., Singleton 4.1943 (MEL); Cumberland Creek, Woods Pt. Rd., 
Marysville, on rocks and fallen logs in bed of creek, Singleton 4.1943 (MEL). 


Tasmania: North: Mt. Barrow, 2,400 ft. alt. on trunks of Nothofagus cunninghamii 
Tindale 12.1954 (NSW P6998); ditto, 2,700 ft. alt., on fallen logs of N. cunninghamii, Burns’ 
6.1962 (NSW P8389); ditto, 3,300 ft. alt., on fallen logs of N. cunninghamii , in deep shade, 
eastern aspect. Burns 6.1962 (NSW P8388); head of Quamby's Bluff, Meander River, Stuart 
1.1849 (MEL). North-West: Cradle Mountain, Sutton 1.1911 (MEL); Henty River, 
Queenstown to Zeehan, in mixed Nothofagiis forest, Vickery 1.1949 (NSW P4990); Franklin 
River (Lyell Highway), in mixed Nothofagus forest, Johnson 1.1949 (NSW P4992, P4991); The 
Acheron (Rivulet, a tributary of the Jane River before it joins the Franklin River), Gunn No. 1511, 
2.1845 (NSW P5066). East: South Esk River, Herb. Mueller (MEL); South Esk, shady 
creeks amongst moss. No. 492 (MEL); St. Patrick's River, Gunn No. 1511,4.1845 (NSW P5064). 
South-East: Uxbridge, Rodway (HO 5042); near the junction of the Huon and Picton Rivers, 
epiphytic on a fallen tree in river brush, Whaite No. 2170, 1.1962 (NSW P8361); Picton River, 
Johnston (MEL); Lady’s Bay, near South Port, Stuart No. 1533, 4.1856 (MEL); South Port, 
Stuart No. 1533, 4.1856 (MEL). South-West: Port Davey Track, Mt. Mueller, on mossy 
log in myrtle forest, 2,500 ft. alt., Olsen 1.1941 (HO 5043); Mt. Mueller, near tributary of the 
Weld River, Olsen 1.1941 (HO 5048). 

Absolute Synonym: Trichomanes peltatum Poir. in Lam., Encyc. VIII (1808) 76. 

Equivalent Synonyms: Hymenophyllum unilaterale Willd., Sp. PI. V (1810) 521, Isotype: 
lie de Mascareigne, an X, £t Willdenow sous le no. 117 (P), examined. Hymenophyllum meyeri 
Presl, Hymen. (1843?) 50, Isotype?: Capite bonae spei (Cape of Good Hope), Drege (L 
910.28_46). 

H. peltatum was omitted in Moore & Betche, Handb. FI. N.S.W. (1893) and Maiden, 
& Betche, Census N.S.W. PI. (1916), being recorded from the Australian mainland for the first 
time by Watts in Proc. Linn. Soc. XLI (1916) 381. 

The specimens of H. peltatum from the mainland do not differ morphologically from 
those collected in the State of Tasmania. Specimens from the latter island (e.g. NSW P4990 
and P6998) have pitted laminal cell walls just as in the material from Barrington Tops, N.S.W. 
(e.g. NSW PI234) and the Thredbo River Gorge (e.g. NSW P7213). 

In New South Wales this species is uncommon, being restricted to the head of the Bellenger 
River, the Barrington Tops district and the Southern Alps where it occurs at high altitudes. 


16 


201 . Hymenophyllaceae. ( Tindale ) 


H. peltatum is more common in Tasmania where it is found on the mountainsides in Nothofagus 
cunninghamii forest as well as in mixed Nothofagus forest occurring along rivers. The latter 
forest consists of N. cunninghamii, Atherosperma, Anopterus, Eueryphia and Anodopetalum. 
//. peltatum occurs mainly on fallen logs of N. cunninghamii but occasionally on living trees of 
this species. It also occurs on the mossy boles of Atherosperma moschatum and on rock faces 
especially in Victoria, being found on some of the higher mountains listed above, as well as on 
Mt. Ellery, East Gippsland, where it was collected by Willis on steep granitic slopes at about 
3,600 ft. alt. in December, 1951 (MEL). In addition' to the localities cited above, H. peltatum 
was collected at the head of the Royston River by Willis in 1943. 

A chromosome count of 2n = 22 was obtained by the author from material collected 
at Mt. Barrow, near Launceston, Tasmania, by T. Burns in June, 1962 (NSW P8388), (No. 201, 
Plate VII). I wish to thank Mr. Burns for sending these plants, also Dr. Barbara Briggs who 
checked the count and Mr. J. Peacock who photographed the cell. Brownlie published a 
chromosome count of n = 11 for material of H. peltatum collected in New Zealand (see Trans. 
Roy. Soc. N.Z. LXXXV (1958) 213). The material from New Zealand which 1 have examined 
is morphologically very similar to Australian specimens of this species. 

The primary pinnae of the 3 Mascarene Island specimens of H. peltatum which were 
kindly sent on loan by the Director, Museum d'Histoire Naturelle, Paris, are more closely spaced 
than in the Australian and New Zealand material, also 6 to 8 lobes are quite common on the 
primary pinnae in material collected in the Mascarene Islands, whereas 2 to 4 or rarely 6 lobes 
are found in the Australian and New Zealand specimens. There appears to be no difference 
in the setting, spacing or type of teeth on the fronds. 

Dr. E. A. Schelpe very kindly sent me a specimen of H. peltatum which he collected on 
Table Mountain, Cape Peninsula, South Africa, in April, 1954 (Schelpe No. 4652). This material 
is a good match for Australian specimens of this species. 

HymenophyUum wilsonii Hook, is a closely allied species which was formerly included 
in It. peltatum by a number of authors, but the chromosome number is 2n = 36 (see Clapham, 
Tutin & Warburg, FI. Brit. Isles (1952) 17). H. wilsonii occurs in Great Britain, Ireland, France, 
Norway and Macaronesia. 


3. HymenophyUum cupressiforme Labill., Nov. Holl. PI. Sp. II (1806) 102, 
t. 250, f. 2; Copel. in Phil. Journ. Sci. LXIV (1937) 91; Wakefield in Viet. Nat. 
LVIII (1941) 110; Wakefield, Ferns of Victoria and Tasmania (1955) 6, 4 f. 5, 7 
photo. 


Habitat in mountain and coastal gullies, mostly in subtropical rainforests 
but also in Nothofagus cunninghamii and N. moorei temperate rainforests, rarely in 
rocky gullies dominated by dry sclerophyll forests, usually forming large, matted 
patches on moist rock faces near creeks or waterfalls, under dripping rock ledges 
or in shaded recesses of sandstone cliffs, more uncommonly in mine-shafts or 
epiphytic on trees or trunks of Todea barbara, occurring in sandstone, granitic, 
basaltic or limestone country. Rhizome filiform, wiry, long-creeping, extensively 
branched, dark brown, 0-1 to 0-3 mm. in diam., sparsely ferruginous-tomentose 
on some of the younger parts. Stipes distant, 0-6 to 3-5 cm. long, 0-1 to 
0-3 mm. broad near the middle, dark brown, glabrous, dull or slightly glossy, smooth, 
almost imperceptibly winged near the apex or sometimes for f of the length of the 
stipe or rarely almost to the base (e.g. NSW P2047), clothed at the base with a few, 
dark reddish, 3- or 4-celled, unbranched hairs. Main rhachis with a very narrow, 
entire or toothed wing which becomes almost imperceptible near the base, dull, 
smooth, glabrous or sometimes bearing a few, undivided, 5- or 6-celled hairs with 
dark red septae. Fronds 2 to 9 cm. long including the stipes, often decurved, dark 
green, erect. Lamina 1-5 to 6-2 cm. long, 1-5 to 2-5 cm. broad, 2-pinnate-pinnatifid, 
usually elliptical-oblong, the apex rounded or subacute, tapering towards the base, 
the lowest 2 or 3 pairs of primary pinnae often half as long as the middle pinnae. 
Primary pinnae with a short winged pedicel except in the basal pairs where the pedicel 
is unwinged, flabellate-rhomboidal, alternate, remote to slightly crowded, usually 


201 . Hymenophyllaceae. ( Tindale ) 


17 


branched on the upper side but lobes do occur on the lower side, the apex broadly 
rounded. Secondary pinnae sessile, simple or bifid, finger-like. Ultimate segments 
1-5 to 5 mm. long, 0-5 to 1 mm. broad, cultrate (6: 1) to narrowly oblong (3: 1), 
the margin with straight or rarely curved teeth, the apex toothed and truncate or 
sometimes obtuse. Veins free, with a branch in each lobe, not reaching the apex 
of each ultimate segment, glabrous. Sori indusiate, marginal, each borne on a 
short, lateral segment, sometimes with a basiscopic subtending segment, usually 
solitary but up to 6 on a primary pinna in rare cases, mostly borne close to the main 
rhachis on the acroscopic side of the secondary rhachises but occasionally on the 
basiscopic side too, occurring in the upper part of the lamina, sunken or free. 
Involucre 1 to 1-5 mm. long, 1-3 to 2 mm. broad at maturity, obovate, 2-valved, 
with a short tube, the valves suborbicular, divided down to the cuneate base, much 
wider than the ultimate segments, persistent, green, the margin almost entire to 
erose or denticulate, glabrous or rarely with a few, fawn, clavate hairs near the base 
of the involucre. Receptacle included or slightly exserted at maturity, club-shaped, 
up to 0-8 mm. broad at the apex. Sporangia subsessile, numerous (often about 40), 
bright orange-red when mature, the cluster of sporangia up to 1-8 mm. in diam., 
each sporangium turbinate, the obliquely horizontal annulus composed of 17 to 19 
indurated cells and 7 to 9 thin-walled cells. Spores subtriangular to globose, 
trilete, greenish, subgranulate, with long narrow laesurae, equat. diam. 41-25 to 
52-5[j. x polar diam. 27-5 to 48-75pt. (No. 201, PI. II, f. 4.) 

Distribution: Australia (south-eastern Queensland, New South Wales, Victoria and 
Tasmania). 

Holotype: Nova Hollandia et Terra Diemen, Herb. Webbianum, ex Herb. Labillardiere 
(FLOR), examined. Labillardiere’s original description in his handwriting is affixed to this 
specimen. The valves of the involucres are denticulate at the broadly rounded apices but the 
sides of the lobes are entire. NSW PI 170 collected at Launceston, South Esk, Tasmania, by 
Lucas in May, 1924, is a good match for the holotype. 

Queensland: Lamington National Park, Elabana Falls track, alt. ca. 2,800 ft., on rock 
in rainforest. Smith & Tindale 8.1956 (NSW P7458); Lamington National Park, in rainforest 
near the head of Tooloona Creek, alt. ca. 3,400 ft., epiphytic fern. Smith No. 10912, 3.1960 
(NSW P8411); Binna Burra, Lamington National Park, Floyd 5.1961 (Forestry Office, Coff’s 
Harbour). 

New South Wales: N. Tablelands: Point Lookout, in Nothofagus moorei association, 
Davis 7.1956 (NSW P8355); ditto, 5,000 ft. alt., on rocks with mosses, Davis 7.1956 (NSW 
P7754). North Coast: Mt. Warning, about 2,000 ft. alt.. Rodway No. 10886, 10.1939 
(specimen sent to Kew No. 2979) (NSW P8415); Dorrigo, Boorman 3.1909 (NSW P2073); 
The Dome, Dorrigo, in subtropical rainforest on rocks, Tindale 4.1944 (NSW P1605, P1988); 
The Rapids, Ellenborough River, Watts 4.1915 (NSW PI 105); Hell's Delight, Barrington, 
Fraser & Vickery 1.1934 (NSW P8416). Central Western Slopes: Munghorn Hill, Mudgee- 
Wollar Rd., alt. 1,875 ft., on a moist rock-face, rocky gully in dry sclerophyll forest, sandstone, 
Johnson & Constable 8.1950 (NSW P6284). Central Tablelands: Natural Bridge Creek, 
Mt. Cameron, on rock in bushy gully. Constable 12.1948 (NSW P5307; U; BM; L); Bowen's 
Creek, Bilpin-Mt. Irvine, alt. 1,850 ft., thickly growing on moist rock face in rainforest gully, 
basalt, Tindale & Constable 3.1952 (NSW P2047; US; K); Waterfall Creek, Mt. Wilson, 
alt. 3,000 ft., on a rock in rainforest, basalt, Melville & Tindale 4.1953 (NSW P6404); Waterfall 
Gully, Mt. Wilson, growing on tree in rainforest, Melville & Tindale 4.1953 (NSW P6406; L; 
US); Happy Valley, Mt. Wilson, Watts 12.1914 (NSW PI 117); Grose Valley, Watts 8.1914 
(NSW PI 114); Tomah Creek, Mt. Tomah, in rainforest, Ingram 5.1956 (NSW P7470); Mill 
Creek, Mt. Tomah, on rocks. Ingram 9.1956 (NSW P8419); South Kurrajong, Stephenson 
7.1892 (NSW P2076); Mt. Victoria, Watts 8.1914 (NSW PI 106); Erskine Creek, Kings Tableland, 
18 miles SE of Wentworth Falls, frequent on rocks in rainforest gully. Constable 5.1956 (NSW 
P7413); Horseshoe Falls, Blackheath, on flat rock, sandstone, Constable 1.1949 (NSW P5814); 
Govett’s Leap, Blackheath, 3,200 ft. alt., on moist rocks at bottom of falls. Constable 3.1957 
(NSW P7890); Wentworth Falls, on moist rock face, sandstone. Constable 10.1948 (NSW 
P5640); Valley of the Waters, Watts 12.1914 (NSW PI 108-9); Victor Falls, Hazelbrook, alt. 
1,750 ft., on damp rocks, sandstone country, Constable 3.1949 (NSW P5604); Hazelbrook, 
Maiden 5.1905 (NSW P2079); The Lake, Hazelbrook, 2,000 ft. alt., on moist rocks near as 


18 


201 . Hymenophyllaceae. ( Tindale) 


waterfall, sandstone. Constable 3.1949 (NSW P6247; US; K; BM); head of Linden Creek, 
Linden, 1,900 ft. alt., on moist rocks in creek, frequent, sandstone. Constable 1.1954 (NSW 
P6725; UC; KYO); Springwood. alt. 750 ft., on sloping rock in gully, sandstone country. 
Constable 2.1949 (NSW P5605); ditto. Watts 12.1914 (NSW PI 113); Springwood South, on 
rock face in creek, sandstone country. Constable 2.1949 (NSW P5603); Mittagong, Dixon 
1882 (MEL); Belmore Falls, near Robertson, Rodway No. 10870, 10.1932? (NSW P8418). 
Central Coast: Newport, Watts 8.1915 (NSW PI 110, P2074); Narrabeen, on rocks on top of 
a waterfall, Mair & Tindale 5.1949 (NSW P5594); junction of Rocky Creek and Middle Harbour, 
3 miles E of Pymble, alt. 10 ft., on moist sandstone rock face in gully, Johnson 10.1960 (NSW 
P8403); Port Jackson, Siegert 1884 (MEL); Newtown, Collie 1882 (MEL); Lucas Heights, 
near Menai, in gully near creek in shaded situation on rocks, Mair 3.1955 (NSW P8404); National 
Park, Boorman 7.1904 (NSW P2081); ditto, on rocks beside creek near rainforest, Tindale 

2.1944 (NSW P43); near Waterfall, National Park, Rodway 7.1933 (specimen sent to Kew No. 
1147) (NSW P8412); Loddon Falls, W of Bulli Pass, on rock face on W side of falls, Evans 
5.1961 (NSW P8405; MEL); Austinmer, Rodway 4.1934 (specimen sent to Kew No. 1387) 
(NSW P8409); Mt. Kembla, Fletcher 11.1891 (NSW P1168, PI 116); Minnamurra Falls, 3 miles 
W of Jambcroo, 1,200 ft. alt., in rainforest on boulder, in mist region, Judd 5.1954 (NSW P6789); 
ditto, on face of boulder in creek near the falls, Tindale 4.1953 (NSW P6688); ditto, frequent 
on moist rocks, in dense rainforest near the falls, alt. 100 ft., Constable 2.1959 (NSW P8306); 
Woodhill near Berry, on sandstone boulder, on top of range, 1,000 ft. alt., Rodway No. 10878, 
4.1930 (NSW P8406); summit of Cambewarra Range, on rock, 1,500 ft. alt.. Rodway No. 10875, 
6.1929 (NSW P8407); Cambewarra Mt., Nowra, Boorman 2.1910 (NSW P2075); Paradise 
Gully, Kangaroo Valley, on a boulder in a shady creek. Rodway No. 10877, 1.1939 (NSW P8408); 
Bomaderry Creek, sea level, on a sandstone boulder, Rodway No. 10874, 10.1930 (specimen 
sent to Kew No. 105) (NSW P8402). South Coast: The Falls, Nowra, Watts 5.1903 (NSW 
PI 112); summit of Pigeon House Mountain, near Milton, in cracks of sandstone cliffs, Rodway 
No. 10880, 11.1932? (NSW P8420); Bateman’s Bay, Boorman 6.1906 (NSW P2072); NE slopes 
of Mt. Dromedary, Central Tilba, 1,400 ft. alt., abundant on rock face in rainforest. Constable 
9.1953 (NSW P7143); near Mt. Dromedary, Reader 1880 (MEL); Twofold Bay, White 1881 
(MEL); Disaster Bay, Green Cape, Rodway (NSW P8421); Murrica River (ca. W of Disaster 
Bay), in rainforest gully on rocks, alt. 15 m., with Piychomnion aciculare, Constable 10.1954 
(NSW P7899). Southern Tablelands: Sugar Loaf Mountain, Braidwood, Bauerlen 8.1898 
(NSW P2083). 

Victoria: Snowy Creek, Ovens River, NE Victoria, McCann 1881 (MEL); Buffalo 
Ranges, on moist shaded rocks, as well as on stems of Todea barbara, Mueller 3.1853 (MEL); 
Dandongadale Falls, Mt. Cobbler Plateau, at about 4,000 ft. alt., on dripping rock ledges, Willis 

1.1945 (MEL); near Mt. Ellery, Merrah 1887 (MEL); Cape Howe, Walter (MEL); between 
Bemm and Coombinbah Rivers, E Gippsland, Sayer 1887 (MEL); near Lakes Entrance, ferntree 
gully, on decaying logs of lilly pilly, 100 ft. alt., Stirling 5.1882 (MEL); Acheron Way, between 
Marysville and Warburton, at about 3,000 ft. alt., Tadgell (MEL); Blacks’ Spur, Fcrnshaw, 
Walter 4.1872 (MEL); Warburton, Walts 1.1919 (NSW PI 126); Mt. Juliet, (near Burgoyne), 
Sullivan 3.1872 (MEL); Walhalla, Herb. Mueller (MEL); Clematis Gully, Kallista, Singleton 
1943 (MEL); near the head of Hardy’s Gully, Kallista, Dandenong Ranges, Willis 1935 (MEL); 
Dandenong Ranges, on tree-fern stems, Mueller 1.1853 (MEL); Mt. Dandenong, Cowle (MEL); 
Olinda Creek (cascades), Dandenong Range, Singleton 5.1943 (MEL); Mirboo North, Howitt 
No. 963 (MEL); Art.air's Seat, Mornington Peninsula, Baldry 9.1883 (MEL); Sealers Cove, 
on stems of ferntrees (MEL); Creswick. in mine shafts, uncommon, Willis 11.1931 (MEL); 
Stony Rises, W of Colac, Singleton 8.1944 (MEL); Apollo Bay, Marriner? (MEL); Apollo 
Bay. Mueller (MEL); Mt. Cole, probably the granitic fern gullies at Middle Creek Falls, Green 
(MEL); Mackey’s Peak, at Hall’s Gap, Grampians, in damp shaded recesses on sandstone 
cliffs, at ca. 1,850 ft. alt., Willis 10.1959 (MEL); Darlot’s Creek, near Portland, Ailitt (MEL). 

Tasmania: North-East: Notley Fern Gorge, near Launceston, 800 ft. alt., epiphytic 
on tree trunk in rainforest gully, Gordon limestone, Tindale 12.1954 (NSW P7077); Distillery 
Creek, near Launceston, common, Gunn No. 1510 (NSW PI 815); ditto, Hannaford 1.1862 
(MEL); Launceston, South Esk, Lucas 5.1924 (NSW PI 170), agrees well with the holotype; 
Mt. Barrow, Dorset, Rupp 1.1922 (NSW P5063); Mt. Victoria, head of Ringarooma River, 
Glover 1883 (MEL); St. Patrick’s River, Gunn No. 1510, 4.1845 (NSW P5062). South-East: 
Derwent (River), Brown Iter Austr. No. 100, 1802-05 (MEL); i mile below Riveaux Falls, 
Huon River, growing with Ctenopteris heterophylla, epiphyte on fallen tree trunk in river bush, 
Whaite 1.1962 (NSW P8422). 

Equivalent Synonym: Hymenophyllum antarcticum Presl, Hymen. (1843) 50, t. XIIA; 
Copel. in Phil. Journ. Sci. LXIV (1937) 81, pi. 40, Syntypes: Synops. Filicum, N. Hollandia, 
Sieber No. 134 (E), examined; Nov. Hollandia, Sieber, Synops. Filicum No. 134 (L 908, 
282 .... 203) (L), examined, according to a note by Copeland part of this specimen was drawn 
by Borbe for plate 40 in Phil. Journ. Sci. LXIV (1937); ditto (W), examined. 


201 . Hymenophyllaceae. {Tin dale) 


19 


Hymenophyllum tunbridgense auctt. non (L.) Sm. has been misapplied to H cupressiforme 
by the following authors:—R. Br„ Prodr. FI. N. Holl. (1810) 159; Benth FI Austr^VII /l8781 
70 .6; F- M. Bail., Syn Queensl FI (1883) 688; F. M. Bail., Lith. Ferns Queensl! (1892) 30 
middle illustration; Moore & Betche, Handb. FI. N.S.W. (1893) 504- F M Bail Oueensl’ 
FI. VI (1902) 1946; Rod way Tasm. FI. (1903) 290; Ewart, FI. Viet. (1931) 32 MeWaTne in 
Proc. Linn. Soc. N.S.W. LXI (1936) 114. K ' ’ Melvaine ln 

I have been unable to retain Hymenophyllum antarciicum Presl as a distinct snecies 
since it is connected to H. cupressiforme by a series of intermediates. The illustration of 
H. antarciicum in Presl's Hymen. (1843) t. XIIA shows an almost entire involucre which is 
characteristic of the syntypes. Typical H. antarticum represents the long, narrow form with 
comparatively short, lower primary pinnae and an involucre which is nearly entire There is 
every variation from the latter type of involucre, c.g. NSW P8405 from Loddon Falls NSW 
to those with a markedly toothed margin, e.g. N.S.W. P7458 from Lamington National' Park’ 
Queensland; SE of Tenterficld, Stuart 11.1874 (MEL) and J mile below Riveaux Falls Tasmania 
(NSW P8422). The last-mentioned specimen is a particularly robust form of H cupressiforme 
the fronds being up to 9 cm. in length including the stipes. However, this species is usually 
characterized by short, broad fronds from 2 to 6 cm. long including the stipes. In some cases 
very depauperate specimens bear sori, e.g. Grass Tree Hill, Tasmania, 1,200 ft. alt. on wet 
sandstone, prostrate, collected by Long in September, 1931 (HO). 


H. cupressiforme is by far the commonest member of the Hymenophyllaceae in south¬ 
eastern Australia where it is found on moist rock-faces in most of the subtropical and temperate 
rainforests on the coast and tablelands. It is less commonly epiphytic on the trunks of trees 
or Todea barbara. In the Munghorn-Wollar district of the Western Slopes, N.S W 
H. cupressiforme is found in rocky gullies in dry sclerophyll forests but this is unusual as it is 
essentially a rainforest species, although it is comparatively hardy. This species does not extend 
to Central or North Queensland. 


H. revolutum Colenso is closely related to H. cupressiforme but the fronds of the former 
species are paler green, more pellucid and more delicate than those of the latter species also the 
involucres of H. revolutum are almost always more sharply toothed than in the other species 
In H. revolutum the marginal teeth of the laminal segments are often falcate although some arc 
straight, whereas in H. cupressiforme they are mostly straight. The main rhachis of H. revolutum 
is only winged towards the apex and the stipe is unwinged, whereas the main rhachis is always 
winged to the base in H. cupressiforme and part of the stipe at least is bordered by a very narrow 
wing. The former species is endemic to New Zealand whereas the latter is restricted to south¬ 
eastern Australia. 


4B. HYMENOPHYLLUM Sm. SUBGENUS CRASPEDOPHYLLUM (Presl) 

Copel. 

Hymenophyllum Sm. sect. Craspedophyllum Presl, Hymen. (1843) 125; subgen. 
Craspedophyllum (Presl) Copel. in Phil. Journ. Sci. LXIV (1937) 163. 

Craspedophyllum (Presl) Copel. in Phil. Journ. Sci. LXVII (1938) 27; Copel Gen 
Fil. (1947) 33. 

Pachyloma van den Bosch, Eerste Bijdrage (1861) 318, non DC. (1828). 

Tiny mat-forming ferns. Rhizome long-creeping, filamentous. Lamina 
0-4 to 3-5 cm. long, either simple or once to thrice forked, the marginal cells usually 
black, the margin glabrous, entire or toothed. Sori terminal. Involucre 2-valved, 
cleft to the base, oval or orbicular, sometimes with a short tube, green or blackish- 
brown, with a border of black cells. Receptacle included, cylindrical. Spores 
tetrahedral-globose to globose. 

Type of Genus: Hymenophyllum marginatum Hook, et Grev. of Australia 
(Queensland, New South Wales and Tasmania). 

There are 2 species in this subgenus of Hymenophyllum, the other species 
being H. armstrongii Bak. which is confined to New Zealand. In Trans. N.Z. Inst. 


20 


201 . Hymenophyllaceae. ( Tindale ) 


LIV (1923) 592, J. E. Holloway pointed out that Hymenophyllum cheesemanii Bak. 
was not worthy of specific rank, since the thickening of the margin had not proved 
a reliable distinction. Giesenhagen in Die Hymenophyllaceen, in Flora LXXIII 
(1890) 441, suggested that the marginal thickening of the lamina kept the margin 
flat and thus protected it from tearing or curling. Another 2 species which may 
belong in this subgenus are Hymenophyllum johorense Holttum (Malaya, Borneo 
and the Philippines) and H. reductum Copel. (the Philippines). Both H. marginatum 
and H. armstongii have a 2-valved, oval or orbicular involucre which is cleft to the 
base and sometimes has a very short tube, whereas the illustrations of H. johorense 
and H. reductum in Phil. Journ. Sci. LXIV (1937) pi. 19-20, show the involucres to 
be of the Meringium-type, i.e. with a hollow, conical tube at the base. According 
to Copeland (l.c. 52) the dark colour in the involucre of H. johorense spreads along 
the whole margin from the hair-bearing teeth and, when the sporangia are mature, 
the margin of the lips of the involucre is dark to a depth of usually 4 cells. In 
Ferns of Malaya (1954) 80, Holttum has pointed out that the margins of the laminas 
in H. armstrongii, H. johorense and H. reductum are characterized by stiff marginal 
hairs consisting of 2 or 3 elongated cells. However, these hairs are absent in 
H. marginatum. Holttum suggested that the 3 former species belong to a much 
reduced group related to Meringium. In his latest paper on this family, Copeland 
in Fern Flora Philippines I (1958) 64, referred both H. johorense and H. reductum 
to Meringium but said that their affinities are uncertain and that they may both 
be better regarded as Microtrichomanes. Cytological evidence may later throw 
some light on this problem. 


Hymenophyllum marginatum Hook, et Grev., Ic. Fil. I (1827) t. 34; Benth., 
FI. Austr. VII (1878) 705; Moore & Betche, Handb. FI. N.S.W. (1893) 504; Rodway, 
Tasm. FI. (1903) 290; C. Chr., Ind. Fil. (1905) 364; Maiden & Betche, Census 
N.S.W. Pi. (1916) 1; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114; Copel. 
in Phil. Journ. Sci. LXIV (1937) 163, pi. 89. 

Habitat in subtropical and temperate rainforests as well as in horizontal 
scrub, often near creeks or waterfalls, forming matted patches on rocks, logs or 
wet cliff-faces. Rhizome long-creeping, branched, smooth, black or very dark 
brown, glossy, non-glaucous, cylindrical, filiform, wiry, sparsely pilose with a few, 
simple, linear (12: 1 or more), red or brown hairs. Stipes very narrowly winged 
at the apex, filiform, 0-3 to 8 mm. long, distant on the rhizome, cylindrical, wiry, 
black or very dark brown, glossy, smooth, clothed near the base with a tuft of simple, 
linear (12: 1 or more), red hairs and near the middle and apex with red tubular 
hairs. Lamina simple, once or more rarely twice or thrice forked, thin, light green 
when dried, darker when fresh, linear (12: 1 or more) to narrowly oblong (3: 1), 
4 to 35 mm. long, 1 to 2-5 mm. broad, glabrous, the older internal cells with wavy- 
thickened walls; margin entire, undulate, glabrous, never toothed, composed of 

1 or 2 rows of glossy, black, obliquely-placed cells; apex of the sterile lobes rounded, 
truncate or emarginate but more or less constricted in the fertile lobes; base gradually 
tapering into the stipes. Midrib black, glossy, without lateral branchlets except 
into the lobes, clothed with a few, reddish, tubular hairs each of which has a short, 
globose, basal cell. Sori indusiate, marginal, solitary at the apex of the frond or 
its lobes. Involucre 0-8 to 1 -5 mm. long, orbicular or oval, valvate to the base or 
sometimes with a very short tube, the cell-walls much pitted, light green, always 
with a marginal band of black, glossy, obliquely-placed cells, the border usually 

2 or 3 cells wide; margin entire, glabrous; base constricted, often clothed with 
similar hairs to those on the midrib. Receptacle included, cylindrical. Sporangia 
about 6 to 10, compressed, orbicular, slightly turbinate, sessile or almost so; annulus 
almost transverse, composed of about 18 to 20 indurated cells and several thin-walled 


201 . Hymenophyllaceae. ( Tindale ) 


21 


cells, dehiscing by a long lateral slit; stomium not a definite group of cells. Spores 
tetrahedral-globose to globose, trilete, white (later green), coarsely granulate to 
subpapillate, equat. diam. 32-81 to 56-25g x polar diam. 30 to 46-88jj.. 

Distribution: Australia (south-eastern Queensland, New South Wales (mainly in the 
Blue Mountains) and Tasmania). There is no record of this fern occurring in Victoria. In 
Proc. Linn. Soc. N.S.W. VIII (1884) 469, there is a note mentioning the collection of this species 
at Milton, South Coast, N.S.W., by W. Bauerlen in 1884. This was probably the first record of 
H. marginatum being collected in New South Wales since Fraser’s original specimens were 
obtained. 

Holotype by Monotypy: Hooker and Greville in Ic. Fil. I (1827) t. 34, cited the type 
locality as “in Nova Hollandia, prope Port Jackson, inter Muscos, Fraser”. There is a 
specimen in the Kew Herbarium labelled “ among mosses, N. Holland, Fraser ” which must 
represent Hooker and Greville’s holotype (examined). Hooker labelled this specimen 
Hymenophyllum marginatum Hook, et Grev., whereas the isotype in the Herbarium of the Botanic 
Gardens, Edinburgh “ Nova Hollandia, Fraser 1825 ” was not determined by Hooker. Another 
Isotype is “ Port Jackson, Fraser (NSW P2089) 

Queensland: Moreton District, Mt. Barney, mostly growing on boulders and forming 
with mosses a dense mat-like cover; collected along creeks from the saddle between North 
and East Peaks to the main Barney Gorge, alt. ca. 2,500 ft., Smith 6.1940 (BRI 27055); Mt. 
Lindsay, alt. about 4,000 ft., small fern on very wet cliff faces near the summit, Everist No. 1123 
6.1935 (BRI 27054). 

New South Wales: Valley of the Waters, Blue Mountains, on the face of wet boulders. 
Watts 9.1912 (NSW P2087, P2900, P2905; MEL; BRI); National Pass, Wentworth Falls, 
Watts 9.1912 (NSW P2088, P2902); foot of Wentworth Falls, Watts 12.1914 (NSW P2903); 
Horseshoe Falls, Blackheath, alt. 2,150 ft., growing on vertical rock face near the falls, sandstone 
formation, in rainforest, Constable 1.1949 (NSW P5815; K; BM; U; US; UC; L; MO; 
LIL); ditto, in spray under a cliff, NE of the falls, on rocks, Watts 1.1911 (NSW P2904, P1910); 
ditto, Watts 6.1911 (NSW P2901); Minnamurra Falls Reserve, about 3 miles W of Jamberoo, 
on a rock face in a deep crevice, very close to a watercourse, ca. 2,000 ft. alt., Judd 10.1958 (NSW 
P8147); Cambewarra, Bauerlen (BRI); near Shoalhaven, Biiuerlen 8.1884 (MEL); Broger’s 
Creek, on moist rocks, Bauerlen 1884 (MEL). 

Tasmania: West: Cradle Mountain, Rodway No. 10894, 12.1915 (NSW P8276); 
Wanderer River, West Coast, Rodway 7.1907 (NSW P2090); Port Davey, Melaleuca Inlet, 
Celery Top Islands’ Bay, matted creeping fern, 1 inch high, on mossy log in horizontal scrub, 
sea level, Olsen 12.1940 (HO H1281), fertile fronds very constricted at the apex; Celery Top 
Island, third island from Narrows in Bathurst Harbour, Port Davey, sea level, Davis No. 1411, 
3.1954 (MEL); Queen’s River, tributary of the King River, West Coast of Tasmania, on trunks 
of Banksia marginata, Moore 1884 (MEL); West Coast, near the Huxley Ranges, Moore 1884 
(MEL). 

Absolute Synonyms: Craspedophyllum marginatum (Hook, et Grev.) Copel. in Phil. 
Journ. Sci. LXVII (1938) 27; Copel., Gen. Fil. (1947) 33; N. A. Wakefield, Ferns of Victoria 
and Tasmania (1955) 4 f. 2, 5. Pachvloma marginatum (Hook, et Grev.) v.d. Bosch in Versl. 
Akad. Wet. Amsterdam XI (1861) 318. 

In south-eastern Queensland H. marginatum has been collected at 2,500 ft. altitude on 
basalt rocks at Mt. Barney in the cloud zone where there would be a considerable amount of 
moisture. It has also been obtained at 4,000 ft. altitude on Mt. Lindsay which is on the border 
between New South Wales and Queensland. In New South Wales this species is mainly found 
at elevations of 2,000 ft. and over in the subtropical rainforests of the Blue Mountains and the 
Minnamurra-Cambcwarra district (near the junction of the Central and South Coast). There 
are no records of this species being collected in Victoria but it is found at Cradle Mountain, 
NW Tasmania, and in the horizontal scrub on the West Coast of Tasmania in the very high 
rainfall area. According to Wakefield in Ferns of Victoria and Tasmania (1955) 5, H. marginatum 
has also been recorded near the Lune River in south-eastern Tasmania. 


4C. HYMENOPHYLLUM Sm. SUBGENUS MECODIUM (Presl ex Copel.) Copel. 

Hymenophyllum Sm. subgenus Mecodium (Presl ex Copel.) Copel. in Phil. 
Journ. Sci. LXIV (1937) 93. Mecodium Presl, Epim. Bot. (1851) 258, nomen; 


22 


201 . Hymenophyllaceae. ( Tindale ) 


Copel. in Phil. Journ. Sci. LXVII (1938) 17; Copel., Gen. Fil. (1947) 33; Atkinson 
in Phytomorphology X (1960) 26-36. 

Diploophyllum van den Bosch, Eerste Bijdage (1861) 322, non Diplophyllum Lehm. 

Epiphytic or rupestral ferns. Rhizome slender, creeping or very rarely short 
and stout. Fronds pinnately compound, the margin entire and glabrous or in 3 
species toothed, the cell-walls mostly thin. Involucre deeply 2-lobed and divided 
to the base, valvate, without a definite tube. Receptacle included. Sporangia 
often spherical. Spores tetrahedral to globose, trilete, the exine thin-walled, 
papillate, granulate or scabrate, often large, sometimes up to 90 ;j. in diam. 
Gametophyte green and ribbon-like, the spore germinating within the sporangium 
first by forming 2 cells with a cross wall, later into 3 cells due to a second wall 
which meets the first at the centre of the spore, so that a triangular plate is formed, 
a cell being cut off at the tip of each of the 3 arms, or in 2 New Zealand species 
(i.e. Hymenophyllum scabrum A. Rich, and H. sanguinolentum (Forst. f.) Sw.) secondary 
walls parallel to the first are formed thus making a 6-celled spherical plate in the 
unopened spore. 

Type of Subgenus: Hymenophyllum sanguinolentum (Forst. f.) Sw. 
(Basionym: Trichomanes sanguinolentum Forst. f.), endemic to New Zealand. 

This is a subgenus of about 100 species many of which occur in the Southern 
Hemisphere. Thirteen species are represented in New Zealand, whereas 8 species 
occur in the tropical, subtropical and temperate rainforests of Australia, 3 of these 
species extend to Tasmania. 

All the Australian species of subgen. Mecodium are characterized by a lamina 
with an entire margin except Hymenophyllum samoense Bak. (Samoa, Fiji to N.E. 
Queensland). In the latter species the margin is obscurely serrulate in some parts 
but mostly entire. H. samoense belongs to a small aberrant group of the subgenus 
Mecodium in which there is a toothed lamina, although otherwise it appears to be 
a typical member of this subgenus. 

In Trans. Roy. Soc. N.Z. LXXXII (1954) 665, Brownlie recorded chromosome 
counts of n = 36 for New Zealand material of Hymenophyllum rarum R. Br. (as 
Mecodium rarum (R. Br.) Copel.) and H. villosum Col. (as Mecodium villosum (Col.) 
Copel.). Later, in Trans. Roy. Soc. N.Z. LXXXV (1958) 213, he also recorded 
counts of n = 36 for the following species collected in New Zealand, viz. H. dilatatum 
(Forst. f.) Sw. (as Mecodium dilatatum (Forst. f.) Copel.), H. scabrum A. Rich, 
(as M. scabrum (A. Rich.) Copel.) and H. demission (Forst. f.) Sw. (as M. demissum 
(Forst. f.) Copel.), as well as n = 72 for H. sanguinolentum (as M. sanguinolentum 
(Forst. f.) Presl) l.c. 215. Later, in Trans. Roy. Soc. N.Z. I (1961) 1 and 2, Brownlie 
obtained a count of n = 36 on larger specimens of the last-mentioned species from 
a different locality in New Zealand. This is an interesting example of polyploidy 
within a species. In this paper he also recorded counts of n = 36 for 2 other New 
Zealand species of this subgenus, viz. H. pulcherrimum Col. (as Mecodium 
pulcherrimum (Col.) Copel.) and H. flexuosum A. Cunn. (as Mecodium flexuosum 
(A. Cunn.) Copel.). A count of n = 36 was recorded by Manton & Sledge in Phil. 
Trans. Roy. Soc. Lond., Ser. B„ Biol. Sci. CCXXXVIII (1954) 144, for material of 
H. flabellatum Labill. collected in New South Wales. Manton & Sledge (l.c. 144) 
made a chromosome count of 2n = 72 (unpaired) for H. javanicum Spreng. which 
extends from Malaya and New Caledonia to North Queensland. 


201 . Hymenophyllaceae. ( Tindale ) 


23 


Although n = 36 is very prevalent amongst the Australian and New Zealand 
species of the subgenus Mecodium, other numbers have been counted in material 
from India, Ceylon and Malaya. In “ Cytology of Hymenophyllaceae ” in Journ 
Genetics LV (1957) 380, a count of n = 21 was made by Mehra and Gurdip Singli 
for material of Hymenophyllum exsertum Wall, ex Hook, collected at Darjeeling, 
India. This number was also recorded by Manton & Sledge in 1954, Lc. 136, for 
specimens of this species collected in Ceylon. The latter cytologists also obtained 
a count of n = 21 for Malayan material of H. serrulatum (Pr.) C. Chr. (l.c. 136). 
Mehra and Gurdip Singh cited a count of n = 27 for specimens of the polymorphic 
species Hymenophyllum polyanthos Sw., collected at Darjeeling, whereas Manton & 
Sledge recorded n = 28 for material of this species forwarded from Ceylon. 
According to Manton & Sledge this also gave clear proof of a monoploid number 
as low as 7 existing in this genus. 

Hymenophyllum whitei Goy, H. walleri Maiden et Betche, H. javanicum, 

H. samoense and H. contiguum (D. A. Smith) Tindale* are found in the tropical 
rainforests of north-eastern Queensland, whereas H. australe Willd. and H. flabellatum 
Labill. range from south-eastern Queensland to Tasmania and H. rarum from the 
Blue Mountains of New South Wales to Tasmania, the 3 latter species being found 
in both subtropical and temperate rainforests. However H. dilatatum which is 
common in New Zealand has been deleted from the list of Australian species, because 
the records are very doubtful. 

I. Stipe and the lower part of the main rhachis not winged. Sori slightly immersed at 

the base. Fronds 2- or 3-pinnate-pinnatitid . h. flabellatum 1. 

1.* Main rhachis winged, the wing usually continuous down the stipe. 

2. Sori free, terminal on the ultimate segments, often in pairs. Stipe broadly 

alate with a crisped or flat wing. Fronds 1- to 3-pinnate-pinnatifid .... H. australe 2. 

2* Sori immersed in the apices of the ultimate segments. Stipe with a short flat 
wing or unwinged. Fronds 1-pinnate-pinnatifid or rarely 2-pinnate-pinnatifid 
. H. rarum 3. 

1. Hymenophyllum flabellatum Labill., N. Holl. PI. Sp. II (1806) 101, pi 250' 
Hook, fil., FI. Tasm. II (1858) 134; Benth., FI. Austr. VII (1878) 705; F. M. Bail.’ 
Syn. Queensl. FI. (1883) 688; F. M. Bail., Lith. Ferns Queensl. (1892) t. 30; Moore 
& Betche, Handb. FI. N.S.W. (1893) 504; F. M. Bail., Queensl. FI. VI (1902) 1946; 
Rodway, Tasm. FI. (1903) 290; Maiden & Betche, Census N.S.W. PI. (1916) 1; 
Ewart, FI. Viet. (1931) 32; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114; 
Copeland in Phil. Journ. Sci. LXIV (1937) 125, pi. 62. 

Habitat in subtropical and temperate rainforests, usually in mountain gullies, 
growing in large matted patches on rocks or as an epiphyte on trees, logs and 
tree-ferns. Rhizome long-creeping, wiry, light brown, 0-3 to 0-8 mm. in diam., 
usually extensively branched and more or less clothed especially at the nodes with 
woolly, pale ferruginous, unbranched hairs several cells in length. Stipes filiform 
or slender, not winged, slightly glossy or dull, dark brown, 1 to 15 cm. long, 0-2 
to 0-5 mm. in diam., glabrescent except for a tuft of woolly hairs at the base. Main 
rhachis sparsely clothed with a few, woolly hairs, winged near the apex of the frond. 
Fronds 5 to 35 cm. long including the stipes, pendulous or the small fronds erect. 
Lamina 2- or 3-pinnate-pinnatifid, pale green or yellowish-green, membranous or 


* Footnote: Hymenophyllum contiguum (D. A. Smith) Tindale comb. nov. (Basionym, Mecodium 
contiguum D. A. Smith in North Queensland Naturalist XIV, No. 80 (1946) 4, 5 (photo) ). 





24 


201 . Hymenophyllaceae. ( Tindale ) 


herbaceous, glabrous or with a few, silky hairs, cultrate (6: 1), narrowly lanceolate 
(6: 1) to narrowly ovate (2: 1) or elliptical (2: 1), in the dwarfed fronds often deltoid, 
ovate (3:2) to orbicular (6: 6), 4 to 20 cm. long, 1-2 to 5 cm. broad; marginal 
cell-walls thin, the internal cell-walls wavy. Primary pinnae flabellate or rhomboidal- 
ovate, deflexed at the apex, acuminate or often produced into a caudate tip, the 
2 or 3 lowest pairs of pinnae distant but the upper pairs overlapping. Lower 
secondary pinnae pseudoflabellate, cuneate at the base. Ultimate segments flat, 
1 to 8 mm. long, 0-5 to 1 mm. broad, cultrate (6: 1), narrowly oblong (3: 1) to 
oblong (2: 1), the margin entire, the apex obtuse or retuse. Veins tree, dichotomous, 
not reaching the margin, often forked near the apex, glabrous. Sori indusiate, 
numerous, 0-5 to 1-5 mm. broad, terminal on the short lateral segments, on all 
parts of the lamina, slightly immersed at the base. Involucre firm, divided to the 
wing into 2 semiorbicular or conical, entire valves. Receptacle included, 
long and slender. Sporangia sessile or subsessile, turbinate, the annulus consisting 
of about 19 or 20 indurated cells and about 8 thin-walled cells. Spores tetrahedral 
to globose, trilete (the laesurae being long ridges), white or greenish, scabrate, 
equat. diam. 35-63 to 45fx X polar diam. 30 to 41-25^. (No. 201, PI. III.) 

Distribution: Australia (south-eastern Queensland, New South Wales, Victoria and 
Tasmania), New Zealand, Tahiti, Samoa, Lord Auckland Islands and possibly Fiji. 

Holotype: Nova Hollandia et Terra Diemen, ex Herb. Labillardit:re, Herb. Webbianum 
(FI), examined. Labillardiere’s original description is affixed to this specimen. The fronds 
of the holotype are rather small, i.e. 3" to 10 cm. long (excluding the stipes) and 2 to 4-2 cm. broad. 
The outer row of laminal cells are not particularly thickened. NSW PI 165 is a good match for 
the holotype except that the rhachises of the former specimen are more pubescent. 

Queensland: Roberts Plateau, Lamington National Park, epiphytic on tree trunks in 
rainforest, fronds very thin, pale green, Smith & Goy No. 73, 1.1938 (BRI); ditto, Shirley (BRI); 
Lamington National Park, ephiphytic on Nothofagus moorei between Bethongabel and Mt. 
Wanungra, alt. ca. 3,800 ft., fronds pendulous, Smith No. 10932, 3.1960 (BRI 25783). 

New South Wales: Head of the Bellenger River, Moore (NSW P2061); William’s River, 
Fraser & Vickery 1.1934 (NSW P2057); Barrington Tops, brush of Nothofagus zone, Fraser & 
Vickery 1.1934 (SYD); Mt. Wilson, Blue Mountains, in rainforest on old log, basalt, Tindale & 
Constable 3.1952 (NSW P2039; K; BM; UC; MO; US; U); Waterfall Gully, Mt. Wilson, 
alt. 3,000 ft., pendulous from a rock ledge near the falls, in rainforest, basalt, Melville & Tindale 

4.1953 (NSW P6399); Happy Valley, Mt. Wilson, Watts 12.1914 (NSW PI 160); Mt. Wilson, 
Brough 7.1926 (SYD); bottom of Govett’s Leap, Blackheath, Watts 4.1903 (NSW PI 161); 
Blackheath, Vickery 9.1936 (NSW P2058); Lawson, Watts 4.1903 (NSW PI 162); Wentworth 
Falls, on damp rocks. Major 4.1933 (SYD); Leura Falls, Stephenson (NSW P2062); Katoomba, 
Forsyth 12.1899 (NSW P1854); Minnamurra Falls, 3 miles W of Jamberoo, 1,200 ft. alt., in 
rainforest on boulder, in mist region, Judd 5.1954 (NSW P6788; UC); Broughton Creek, 
Bauerlen 7.1888 (NSW P2065); Clyde River, Moore (NSW PI 855); Sugar Loaf Mountain, 
Boorman 1.1915 (NSW P2063); ditto, Bauerien 8.1898 (NSW P2052); \ mile NE Trig., Mt. 
Dromedary, Central Tilba, 2,000 ft. alt., plentiful on tree-fern trunks in rainforest, Constable 

9.1953 (NSW P6643; UC; BM); Mt. Dromedary, Tilba Tilba, on trunk of Dicksonia antarctica, 
high on a mountainside, alt. 1,800 ft., Johnson & Constable 9.1960 (NSW P8188; K; BM; 
L; BRI; US; A; U; MO; LIL). 

Victoria: Blacks Spur, Deane 1.1900 (NSW P2059); Acheron Way, treefern gully, 
Melvaine 1.1935 (NSW P1272) and 1.1935 (SYD); Fern-tree Gully, near Melbourne, Nisbet 
(NSW P5061); Kallista, Dandenong Range, in Eucalyptus regnans-Dieksonia-Cyathea forest, 
1,200 ft. alt., epiphyte, rhizome filiform, creeping, fronds light green, Blake No. 7315, 1.1935 
(BRI); Dandenong Ranges, E of Melbourne, pendulous from the branch of a tree in scrub 
along creek. Smith M97, 11.1943 (BRI); Lome, Watts 11.1919 (NSW PI 163); Otway Range, 
Williamson 12.1903 (NSW P2054). 

Tasmania: North-West: Rosebery, Johnson 1.1949 (NSW P5302); Mt. Bischoff, 
Waller No. A14, 1903 (NSW P2053); Strahan, Vickery 1.1949 (NSW P5303). North-East: 
Notley Fern Gorge, near Launceston, on a tree, in a rainforest gully, Gordon limestone, Tindale 
12.1954 (NSW P7079); Mt. Barrow, Co. Dorset, Rupp 1.1922 (NSW P610); Moorina to St. 
Helens, on a tree-fern in beech forest, Ford 9.1950 (NSW P2846); St. Mary’s Pass, Gunn No. 


201 . Hymenophyllaceae. ( Tindale ) 


25 


1514, 2.1846 (NSW P1772). South-East : Cascades Creek, Rupp 9.1920 (NSW P5060); Cascade 
Gully, Hobart, Lucas 5.1924 (NSW PI 165); Eaglehawk Neck, Lucas 11.1924 (NSW P1164); 
Geeveston, Lucas 1.1901 (NSW P2055); Newdegate Caves, Hastings River, epiphytic on heavily 
shaded tree trunk in creek bed, Whaite No. 2210, 1.1961 (NSW P8174); without specific locality, 
Milligan (HO HI275). 

Absolute Synonym: Mecodium flabellatum (Labill.) Copel. in Phil. Journ. Sci. LXVII 
(1938) 21; Wakefield in Viet. Nat. LVII1 (1941) 108; Dobbie & Crookes, New Zealand Ferns 
(1952) 82, 83 (photograph). 

Equivalent Synonym: Hymenophyllum nitens R. Br., Prodr. N. Holl. (1810) 159; 
Hook. & Grev., Ic. Fil. II (1830-1) t. 197, Lectotype: Derwent (Tasmania), R. Brown No. 98, 
Iter Austr., 1802-05 (BM), examined. Isotypes (K.; E), examined. Syntypes: Van Diemen’s 
Land, D. Nelson (BM), examined, and Van Diemen’s Island, R. Brown (K). 

This is a comparatively hardy but very variable species, as plants growing in well-sheltered 
situations in rainforest have narrow, attenuated fronds, whereas those from more exposed 
locations have shorter, more compact laminas. 

H. flabellatum has a wide distribution in south-eastern Australia, New Zealand and 
Polynesia. In Australia it ranges from the mountains of south-eastern Queensland to Tasmania. 
It is relatively common in the subtropical rainforests of Lamington National Park, Queensland, 
as well as in the Blue Mountains and the South Coast of New South Wales. It is also found 
in the Nathofagus moorei temperate rainforests of the McPherson Range on the border of 
Queensland and New South Wales as well as those on the Upper Williams River, N.S.W. In 
addition it is widespread in the Nothofagus cunninghamii temperate rainforests of Victoria and 
Tasmania. This fern occurs at elevations from near sea-level (especially in Tasmania) to 3,800 
it. but it usually favours mountain gullies above 1,800 ft. The slender rhizomes with their 
delicate, pendulous fronds often wind for a yard or more up the trunks of Cyathea spp. or along 
rainforest trees or logs. 

A chromosome count of n = 36 for Hymenophyllum flabellatum was recorded by Manton 
& bledge in Phil. Trans. Roy. Soc. Lond., Ser. B, Biol. Sci. CCXXXV1II (1954) 144. This 
count was made on material collected at Waterfall Creek, Mt. Wilson, Blue Mountains, N.S.W., 

Melville No. 3773 and M. Tindale in April, 1953. In Trans. Roy. Soc. N.Z. LXXXII 
(1954) 665-6, Brownlie also recorded n = 36 for material of this species collected in New Zealand. 


2. Hymenophyllum australe Willd., Sp. PI. V (1810) 527; Maiden & Betche, 
Census N.S.W. PI. (1916) 1; Ewart, FI. Viet. (1931) 32; Melvaine in Proc. Linn. 
Soc. N.S.W., LXI (1936) 114; Copel. in Phil. Journ. Sci. LXIV (1937) 158, pi. 85. 

Habitat in subtropical and temperate rainforests as well as in wet sclerophyll 
forests, rupestral or epiphytic, mostly growing on rocks in matted patches amongst 
mosses or sometimes on the trunks of trees or tree-ferns. Rhizome wiry, brown, 
long-creeping, branched, 0-1 to 1 mm. in diam., clothed with a few, pale ferruginous 
hairs. Stipes wiry, brown, 1 -2 to 10-5 cm. long, 0-3 to 1 mm. broad, aLmost glabrous 
with a few, pale ferruginous hairs near the base, winged almost to the base, with 
the wing plane or slightly crisped. Main rltachis with a comparatively broad, 
plane or slightly crisped wing 1 to 3 mm. wide. Lamina 1- to 3-pinnate-pinnatifid, 
membranous, pale green when young, later dark green, erect or deflexed at the 
apex, narrowly lanceolate (6: 1) to broadly ovate (6: 5) or very narrowly elliptical 
(6: 1) to elliptical (2: 1), 2-5 to 14 cm. long and 1 to 5 cm. broad, the cell-walls 
obscurely pitted. Primary pinnae rhomboidal, 2 to 5 cm. long, spreading, often 
deflexed at the apex, deeply pinnatifid; lower 1 or 2 pairs often slightly shorter 
than the middle pinnae. Secondary pinnae again pinnatifid or irregularly forked. 
Ultimate segments flat or rarely slightly crisped, cultrate (6: 1), the apex obcordate 
or truncate, the margin reinforced, i.e. two cells thick in some places. Veins 
conspicuous, simple or forked, not reaching the apex in the ultimate segments. 
Sori marginal, numerous, 2 sori often on closely proximal lobes, not sunken, terminal 
and solitary on the segments which are rarely constricted below the sori. Involucre 


26 


201 . Hymenophyllaceae. ( Tindale ) 


orbicular, ovate to broadly ovate, mostly broader than the ultimate segments, 
divided to or almost to the base into 2 valves each with an entire or slightly erose 
margin and an acute, broadly rounded, erose or obcordate apex. Receptacle 
enclosed in the early stages, club-shaped. Sporangia subsessile almost spherical 
or slightly turbinate, the annulus large, composed of 21 to 27 indurated cells and 
about 9 or 10 thin-walled cells, the dehiscence being obliquely lateral. Spores 
tetrahedral to almost globose, trilete, greenish, minutely scabrate, equat. diam. 
27-5 to 37-5[i X polar diam. 20-63 to 37-5p.. 

Distribution: New Zealand and Australia (Queensland, New South Wales, Victoria 
and Tasmania). 


Holotype: Nova Hollandia, Labillardicre (Herb. Willdenow No. 20232), (B), two 
photographs examined. 

Queensland: Nerang, Schneider (BRI); Springbrook, on rocks. White 1.1916 (BR1), 
McPherson Range, Try on (BRI). 


New South Wales: Whian Whian State Forest, ca. WNW of Byron Bay, alt. 2,050 ft., 
frequent on moist rocks in rainforest gully, basalt. Constable l - 1953 ,>, 0041 ’ u™' 

William’s River, Colycr 8.1932 (NSW PI270); Gosford, Deane 10.1888 (NSW P3894), Happy 
Valley, Mt. Wilson, Watts & Boorman 5.1915 (NSW P2069, P3838, P3835); ditto, alt., 3, 0 ., 
basalt, growing in a damp cave, Constable 12.1948 (NSW P5642); Mermaid s Cave /xufu/ 
Blackheath, Stephenson (NSW P2066); The Canyon, Blackheath, Melvaine 4.1933 (N8W 
PI268); Valley of the Waters, Watts & Watts 12.1914 (NSW P 3841 ); ditto alt. 1,700 ft., on 
•damp rocks in creek, sandstone country. Constable 2.1949 (NSW P5648); Hippocrenc Falls, 
Rocky Creek, Valley of the Waters, 1,400 ft. alt., sandstone country, Constable 2.194S (INNW 
P5649); Brittania Falls, Wentworth Falls, 1,650 ft. alt., on damp rocks in creek sandstone 
country. Constable 2.1949 (NSW P5651); below Vera Falls, Valley of the Waters Matts 9.1912 
(NSW P3836); Lawson, Watts 4.1903 (NSW P3837); Mt. Kembla, Letcher 11.1891 (NSW 
PI 169); Cambewarra, Bauerlen, 8.1886 (NSW P2800); Conjola, Heron 9.1899 (NSW P2068). 


Victoria: Acheron Way, Melvainc 1.1935 (NSW PI269); Dandenong Ranges, \Valter 
1.1881 (NSW P2067); Kallista, Dandenong Range, in Eucalyptus regnans-Dicksoma-Cyathea 
forest, 1,200 ft. alt., epiphyte, fronds dark green, Blake No. 7314, 1.1935 (BRI); Cement Creek, 
Dandenong Range, in Eucalyptus regnans-Nothofagus forest, amongst moss, fronds light to dark: 
green, 2,200 ft. alt., Blake No. 7247, 1.1935 (BRI); Dandenong Ranges, 20 miles E ol Melbourne, 
Herb. Mueller (MEL); Lome, Watts, 1919 (NSW P3832 and PI 167). 


Tasmania: North-East: Mt. Barrow, Co. Dorset, Rupp 1.1922 (NSW P6792); Notley 
Fern Gorge, near Launceston, 800 ft. all., in rainforest gully, epiphytic on a tree, Gordon limestone, 
Tindale 12.1954 (NSW P7078); ranges near Scottsdale, Lucas 5.1924 (NSW P3892); St. Patrick s 
River, Gunn No. 1513, 10.1845 (NSW P1763). North and West: Hellyer Gorge, Davis 
1.1937 (NSW P5920); Waratah, Lucas 12.1924 (NSW P3833); Gordon River, on tree trunks, 
in dense rainforest, alt. about 30 ft., Johnson 1.1949 (NSW P5618); The Acheron, Gunn No. 
1513, 2.1845 (NSW P1890). South-East: Cascades Creek, Rupp 9.1920 (NSW P4938); Myrtle 
Gully, on rocks and fallen trees, Beadle 1.1935 (NSW P1271); Eaglehawk Neck, Lucas 11.1924 
(NSW P3834); Geeveston, Lucas 1.1901 (NSW P606); Mt, Wellington, 2,000 ft. alt., Rodway 
9.1913 (HO 5053). 

Absolute Synonyms: Sphaerocionium australe (Willd.) Presl, Hymen. (1843) 35. 
Mecodium australe (Willd.) Cope!, in Phil. Journ. Sci. LXVII (1938) 24; Wakefield in Viet. 
Nat. LVIII (1941) 10; Wakefield, Ferns of Victoria & Tasmania (1955) 4 f. 4, 5, photograph 
opposite 15. 

Equivalent Synonyms: Hymenophyllum atrovirens Col. in Tasrn. Journ. Nat. Sci. II 
<1844) 186, Holotype: Waikare Lake, New Zealand, Colenso 12.1841 (WELT), not examined; 
Isotype (K), examined. Hymenophyllum intricatum van den Bosch in Ned. Kruid. Arch. V, 3 
(1863) 168, Holotype; Van Diemen’s Land (i.e. Tasmania), ad rupes juxta fl. St. Patrick, Gunn 
(L910, 38 ... . 102), (L), examined. Mecodium atrovirens (Col.) Copel. in Phil. Journ. Sci. 
LXXIII (1940) 457. 

This species was misidentified as Hymenophyllum javanicum auett. non Spreng., in Benth., 
FI. Austr. VII (1878) 705; Moore & Betche, Handb. Fl. N.S.W. (1893) 504; Rodway, Tasm. 
Fl. (1903) 290. 


201 . Hymenophyllaceae. (Tindale) 


27 


Two photographs of the holotype of Hymenophyllum australe were kindly forwarded to 
me by the courtesy of the Director of the Berlin-Dahlem Herbarium, Germany There are 
2 species represented on this sheet, i.e. a specimen of H. australe and 2 fronds of H. rarum. 
Willdenow's original description referred to the involucres as being ovate, emarcinate at the 
apex and obtusely bidentate which features are characteristic of the fern traditionally known 
as H. australe. However the involucres of H. rarum are entire and acute or rounded at the 
apex but never bidentate. According to van den Bosch’s note on the type sheet, the material 
of H. australe is sterile but presumably Willdenow examined at least one other specimen. Sori 
are present on the material of H. rarum mounted on the type sheet of H. australe. 

There has been a good deal of controversy about whether H. atrovirens of New Zealand 
is identical with H. australe collected in Australia. A specimen of the former collected at Lake 
Ada, Milford Sound, New Zealand, is very similar to NSW P2066 from Mermaid’s Cave, 
Blackheath, N.S.W. 

H. australe is closely related to H. flexuosum A. Cunn. which is a New Zealand species 
with more rounded involucres and very crisped wings on the stipes and rhachiscs. In all the 
specimens of H. flexuosum which I have examined, there is a border of cells one row thick on 
the stipes and rhachises, whereas in H. australe this border is sometimes composed of 2 cells 
in part of the lamina. A chromosome count of n = 36 was recorded for H. flexuosum by 
Brownlie in Trans. Roy. Soc. N.Z. I (1961) 1, 2. 

In H. javanicum Spreng. which has frequently been confused with H. australe , the border 
of the lamina is one cell thick and is composed of oblique (instead of vertical) cells, at least in 
the New Caledonian specimens which I have examined. The segments of the lamina are very 
crisped and the apex of the oblong involucre is lacerated, e.g. in McKee No. 2499 and Vieillard 
No. 2286. Manton & Sledge made a count of 2n = 72 (unpaired) for material of H. javanicum 
collected in Ceylon (see Phil. Trans. Roy. Soc. Lond., Ser. B, Biol. Sci. CCXXXVIII (1954) 
144). On the other hand Mehra & Gurdip Singh published a count of 2n = 108 from material 
of H. javanicum collected at Darjeeling (see Journ. Genetics LV (1957) 391). 

H. australe is a fairly common species in south-eastern Australia especially in the subtropical 
rainforests of the Blue Mountains, N.S.W., as well as in the Nothofagus cunninghamii temperate 
rainforests of Victoria and Tasmania. It sometimes grows with H. flabellatum and Polyphlebium 
venosum on tree trunks. 

Pi nts of H. australe arc usually characterized by a plane wing on the stipes and rhachises. 
Amongst the specimens cited above, only one collection (viz. NSW P3832) from Lome, Victoria, 
has a markedly crisped wing but in NSW P5618 from the Gordon River, Tasmania, the wing is 
slightly crisped. 


3. Hymenophyllum rarum R. Br., Prodr. FI. N. Holl. (1810) 159; Benth., 
FI. Austr. VII (1878) 705; Moore & Betche, Handb. FI. N.S.W. (1893) 504; Rodway, 
Tasm. FI. (1903) 290; Maiden & Betche, Census N.S.W. PI. (1916) 1; Ewart, 
FI. Viet. (1931) 32; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114. 

Habitat subtropical and temperate rainforests, pendulous, forming large, 
matted patches on rocks, tree-ferns ( Dicksortia antarctica) and trees (especially 
Nothofagus cunninghamii and Tristania laurina), often in rocky gorges. Rhizome 
long-creeping, much branched, wiry, black or very dark brown, glossy, cylindrical, 
0-1 to 0-2 mm. in diam., sparsely clothed with hairs which are red-brown, simple 
and septate. Stipes unwinged or sometimes with a very narrow wing almost to 
the base of the stipes, 0-5 to 5 cm. long, 0-1 to 0-4 mm. broad, wiry, filiform, distant 
or often proximal, black or very dark brown, dull or slightly glossy, smooth, glabrous 
except for a few, red-brown, septate, simple hairs at the base. Main rhachis usually 
alate throughout its length with a minute, plane wing. Fronds 1-pinnate-pinnatifid 
or rarely 2-pinnate-pinnatifid, 2 to 14 cm. long including the stipes, with a peculiar 
odour. Lamina 1 -5 to 10-5 cm. long, 1 to 2-5 cm. broad, glabrous, pale green when 
fresh, dark green when dried, membranous, flaccid, irregularly and very narrowly 
elliptical (6: 1) or broadly oblong (3: 2), the internal cells wavy-thickencd especially 
the lumina. Primary pinnae 5 to 20 mm. long, 1 to 9 mm. broad, widely spaced. 


28 


201 . Hymenophyllaceae. ( Tindale ) 


seldom imbricate, rarely simple, mostly 2- to 6-lobed, the lower pairs of pinnae 
shorter and less divided. Secondary pinnae simple or bifid. Ultimate segments 
flat, linear to narrowly oblong, entire, erecto-patent, 1-veined, mostly 3 to 12 mm. 
long, 0-5 to 1-5 mm. broad; apices obtuse or truncate. Veins simple, not reaching 
the apex of each segment. Sori indusiate, marginal, usually 3 to 15 per frond, 
mostly near the apex of the lamina, immersed in the dilated tips of the ultunate 
segments and bordered by conspicuous veinlets. Involucre 1 to 2-2 mm. l°Rg> 
1 to 3 mm. broad, valvate, divided to the middle or lower into 2 lips, broadly 
rhomboidal, cuncate at the base; apex broadly rounded or often pointed; margin 
glabrous, entire or undulate, composed of rectangular cells. Receptacle included, 
very slender. Sporangia subsessile, turbinate, the annulus composed of 16 to 26 
indurated cells and 7 to 11 thin-walled cells. Spores subtriangular to globose, 
trilete, greenish, the laesurae being long, low ridges, the surface subgranulate, equat. 
diam. 33-75 to 75(x x polar diam. 30 to 60a in the material from New South Wales 
and Victoria but 26-25 to 48-75^ X 26-25 to 37-5jx in the specimens from Tasmania. 
(No. 201, PI. II, f. 3.) 

Distribution: Australia (Coast and Tablelands of New South Wales, Victoria and 
Tasmania) as well as New Zealand, the Stewart, Chatham and Auckland Islands. 

Lectotype: Derwent, R. Brown No. 97, Iter Austr., 1802-05 (BM). Isotypes: (Kj 
MEL). Syntype: Van Diemen’s Land, R. Brown (K); Van Diemen’s Island, R. Brown 1804 
(BM; NSW P8426). 

New South Wales: Happy Valley, Mt. Wilson, Watts & Boorman 5.1915 (NSW P612 
PI 125, PI287); ditto, Watts 12.1914 (NSW PI 124); pass above Kiama, Moore 2.1890 (NSW 
P1853; MEL); Minnamurra Falls, 3 miles W of Jamberoo, in a subtropical rainforest, growing 
on the face of a boulder in a ravine, 1,200 ft. alt., Judd 5.1954 (NSW P6785); ditto, on moist 
rocks in deep sheltered rocky gorges, Judd 7.1956 (NSW P7420); ditto, Judd 5.1955 (NSW 
P7165). 

Victoria: Mt. Ellery, East Gippsland, among shaded granite boulders in wet fern gully 
on eastern slopes at about 3,500 ft. alt., Willis 12.1951 (MEL); Mt. Drummer, East ''ippsland, 
fairly abundant on trunks of Tristania and Dicksonia, Wakefield 1.1939 (NSW P611); Pipeclay 
Creek, Orbost, Wakefield 8.1940 (NSW P1284); Cumberland Creek, near Marysville, Singleton 

4.1943 (MEL); Cement Creek, 4 miles N of Warburton-Acheron Way crossing, on Nothofagus 
trunks, Swaby 1.1948 (MEL); Sealers Cove, Wilson’s Promontary, F. Mueller 8.1954 (MEL); 
McCrae Creek, Beenak, about 12 miles NE of Gembrook, on the gnarled mossy buttresses of 
old myrtle beeches (Nothofagus cunninghamii), Willis & Bond 7.1948 (MEL); Melba Gully at 
the head of the Johanna River, Otway Ranges, on tree-fern butts in Nothofagus forest, Singleton 

1.1943 (MEL); Weeaproinah, towards Cape Otway, SW of Melbourne, grows only on Nothofagus , 
Swaby 1.1932 (NSW P1297); Beech Forest, Otways, Swaby 12.1931 (MEL); Apollo Bay, 
Mueller (MEL); Otway Forest, Swaby 1.1931 (MEL). 

Tasmania: North-West: Mt. Bischoff, Waller No. A13, 1903 (NSW P1285); Corinna, 
in rainforest, flood bank. Jackson 1.1954 fHO 5044). North-East: head of the Don River, 
on trunks of tree-ferns, Rupp 1.1915 (MEL); Ranges near Scottsdale, Lucas 5.1924 (NSW 
PI 120); Mt. Victoria, head of Ringarooma River, Glover 1883 (MEL); Mt. Barrow, Co. 
Dorset, Rupp 1.1922 (NSW P5638); ditto, 3,300 ft. alt., on tree-ferns in temperate rainforest, 
Burns 6.1962 (NSW P8435). Central: Gordon Vale Track, 1 mile from the Florentine River, 
epiphytic on tree trunk in beech forest, common, fronds 2-3 inches high, Whaitc No. 2410, 
2.1961 (NSW P8171); River Gordon, Macquarie Harbour, forest, Gunn No. 1512 (NSW 
PI761); west side of Lake St. Clair, growing on face of rock in Fagus (i.e. Nothofagus) forest, 
Gunn No. 1512, 1.1847 (NSW P1819). South-East: Cascade Gully, Hobart, Lucas 5.1924 
(NSW PI 121); Myrtle Gully, Hobart, 1,000 ft. alt., creeping plant, Olsen 9.1936 (HO H1277); 
Collinsvale in myrtle forest, in moist gully, on stem of tree-fern, 1,000 ft. alt., Atkinson 7.1930 
(HO 5046); Newdegate Cave, Hastings River, epiphytic on fallen tree in thick scrub on limestone, 
Whaite No. 2208, 1.1961 (NSW P8173); Bruny Island, between Adventure Bay and Lunawanna, 
1,000 ft. alt., in rainforest on Forestry Roads, climbing up tree-trunks, Phillips 2.1962 (NSW 
P8339). 

Absolute Synonym: Mecodium rarum (R. Br.) Copel. in Phil. Journ. Sci. LXVII (1938) 
21; Crookes & Dobbie, New Zealand Ferns, ed. 5 (1952?) 60, 61 (photo). 


201 . Hymenophyllaceae. (Tin dale) 


29 


Equivalent Synonyms: Hymenophyllum semibivalve Hook, el Grev., le. Fil. 1 (1828) 
t. 83, Holotype: N. Zealand, A. M(enzies), labelled Hym. asplenioides (K), examined. Hooker 
” Greville stated that the specimens collected by Menzies were labelled Hym. asplenioides. 
Lhe fronds on the sheet at Kew Herbarium are not those figured on t. 83 in Hooker & Greville's 
leones Filicum but they belong to the same species. Hymenophyllum imbricatum Col. in Tasm 
Journ. Nat. Sci. 11 (1846) 187, non Blume 1828, Isotype?: A specimen at Kew Herbarium 
labelled by Colenso as “ fol. 5. //. imbricatum Col. haud facile cum H. raro conjungam ". examined. 
Hymenophyllum gunnii Bak. in Hook. & Bak., Syn. Fil. (1874) 463, Holotype: Van Diemen’s 
Land, Gunn (i.e. York Town, R. C. Gunn 23.10.1844), (K), examined. 

In a letter dated 16th February, 1861, written by van den Bosch from Goes, Zeeland, 
Netherlands, to Sir William Hooker (letter No. 374, Vol. XLVlll, Correspondence from France 
1858-65, Germany 1855-61, at Kew), there is the following paragraph “ Among the specimens of 
Hymenophyllum rarum of your Herbarium there occurred specimens gathered by Mr. Gunn 
in van Diemen’s Land. Believing them to be new, I determined them as H. gunnii n. sp. but 
afterwards 1 perceived my mistake and hold them now not different from H. australe W. to which 
belong also the specimens of Menzies in the same envelope.” However, Baker published 
H. gunnii in Hooker & Baker’s Synopsis Filicum, ed. 2 (1874) 463, attributing it to van den 
Bosch. NSW PI 120 collected by Lucas in the Ranges near Scottsdale, Tasmania, is the best 
match for the holotype cited above. 

Some forms of 77. rarum have comparatively undivided fronds, e.g. NSW P1761 collected 
by Gunn at the Gordon River, Tasmania, and NSW PI 125 obtained at Happy Valley, Mt. Wilson, 
N.S.W., by Watts & Boorman. In these specimens a sorus may terminate a simple primary 
pinna especially towards the apex of the lamina, although some fronds in both collections have 
more divided pinnae. Other specimens, e.g. NSW PI 120 from the ranges near Scottsdale, 
Tasmania, have deeply lobed primary pinnae, e.g. up to 9 ultimate segments. The type collections 
of both 77. rarum and 77. gunnii represent the more divided form of this species with narrow 
segments. 

Hymenophyllum imbricatum Col. is a form of 77. rarum front New Zealand but it is 
characterized by very imbricate pinnae. Two syntypes were cited by Colenso, viz. 1) on edges 
of rocky precipices, in dry spots overhanging water, at Pataua, near Wangarei, E. Coast, 1842, 
and 2) on reclining timber, in woods between Wangarei and the Bay of Islands, 1840. No 
locality was cited on the specimen of 77. imbricatum collected by Colenso at Kew Herbarium. 

Hymenophyllum capense Schrad. of South Africa, is very closely allied to the New Zealand 
form of 77. rarum (especially Ruahine Range, New Zealand 11.1919 (BR1 28943) ) which is 
characterized by narrow overlapping segments and closely spaced primary pinnae. 1 wish to 
thank Dr. E. A. Schelpe for the material of 77. capense which he so kindly forwarded. 

In Trans. Roy. Soc. N.Z. LXXX1I (1954) 665, Brownlie recorded a chromosome count 
°f n 36 for material of 77. rarum collected in New Zealand. 


4D. HYMENOPHYLLUM SUBGENUS MERINGIUM (Presl) Copel. 

Hymenophyllum subgenus Meringium (Presl) Copel. in Phil. Journ. Sci. 
LX1V (1937) 14. Meringium Presl, Hymen. (1843) 116, as a genus; Copel. in Phil. 
Journ. Sci. LXVII (1938) 39; Copel., Gen. Fil. (1947) 35. 

Myrmecostylum Presl, Hymen. (1843) 27, pi. 10A. 

Ptychophyllum Presl, Hymen. (1843) 28, pi. ME. 

Mostly small, epiphytic ferns. Rhizome creeping, narrow. Lamina pinnately 
decompound, mostly with thick, pitted cell-walls; margin minutely serrulate or 
rarely entire, never hairy. Axes alate, with a narrow wing 1 cell thick on the minor 
axes or on all the axes, usually hairy on the lower surface. False veins absent. 
Involucre tubular at the base, with 2 large lips. Receptacle slender and usually 
long-exserted (except included or slightly exserted in Hymenophyllum bivalve). 
Sporangia large, sessile, maturing from the top of the receptacle downwards. Spores 
tetrahedral or globose-tetrahedral. 

G 99111 — [ 2 





30 


201 . Hynienopliyllaceae. ( Tindale) 


Type of Subgenus: Hymenophyllum meyenianum (Presl) Copel. (a synonym 
of Hymenophyllum serrulalum (Presl) C. Chr.), a native of the Philippines. Sumatra, 
Malaya to New Guinea. 

This is a comparatively large subgenus of about 60 species some of which 
are found in Malaysia, New Zealand and Australia but other species do occur in 
Formosa, Ceylon, Africa and non-tropical South America. H. kerianwn Watts, 
//. pseudotunbridgense Watts, H. babindae Watts, and //. lobbii Moore arc found in 
the tropical rainforests of north-eastern Queensland, whereas H. bivalve (Forst. f.) 
Sw. occurs in New South Wales and south-eastern Queensland. 

Unlike the other genera and subgenera of the Hymenophyllum- group found 
in south-eastern Australia, subgenus Meringium is characterized by an involucre 
with a well-developed tube. Because of the indefinite receptacle and the form of 
the involucre, subgen. Meringium appears to be more closely related to the 
Trichomanes- group of genera. 

The subgenus Hymenophyllum appears to be joined to subgen. Meringium 
by several dwarf species, viz. H. pumilum Moore (New South Wales), H. moorei 
Bak. (Lord Howe Island), H. pumilio Ros. (New Caledonia) and H. minimum A. 
Rich. (New Zealand). It is difficult to ascertain by the degree of dissection of the 
involucre in which subgenus these tiny species should be placed. 

Very few chromosome counts have been made for members of subgenus 
Meringium but the base numbers so far recorded are 7, 11 and 13. Brownlie made 
a count of n = 22 for Hymenophyllum bivalve (Forst. f.) Sw. (as Meringium bivalve 
(Forst. f.) Copel.) in Trans. Roy. Soc. N.Z. LXXXV (1958) 213. He also recorded 
n = 26 for Hymenophyllum multifidum (Forst. f.) Sw. (syn. Meringium multifidum 
(Forst. f.) Copel.) in Trans. Roy. Soc. N.Z. LXXX11 (1954) 665-6. Manton recorded 
a count of n = 21 for Hymenophyllum serrulalum (Pr.) C. Chr. (syn. Meringium 
meyenianum Presl) in the Appendix to Holttum, Ferns of Malaya (1954) 623. 
A relationship is suggested to several members of subgen. Hymenophyllum , e.g. 
H. pe/tatum (n = 11, Brownlie), H. revolutum Col. (n = 22, Brownlie), II. wilsonii 
(n = 18, Manton) and H. lunbridgense (n = 13, Manton). 

The base numbers 9, 11 and 13 have been recorded for the subgen. 
Hymenophyllum whereas the base number 9 has been consistently found in species 
of the subgen. Mecodium. 


Hymenophyllum bivalve (Forst. f.) Sw. in Schrad. Journ. 11 (1800), 1801, 
99; Swartz. Syn. Fil. (1806) 146, 372; Hook., Sp. Fil. I (1846) 98. pi. 35D; C. Chr., 
Ind. Fil. (1905) 357; Cheeseman, Man. N.Z. FI. (1925) 15; Copeland in Phil. 
Journ. Sci. LXIV (1937) 66, pi. 30. f. 4-6. 

Habitat in temperate or subtropical rainforests either terrestrial in large 
matted patches, growing on boulders in ravines in mist regions or as an epiphyte 
on tree trunks or old logs. Rhizome long-creeping, wiry, branched, 0-3 to 0-8 mm. 
in diam., clothed sparsely especially at the nodes with hairs which are dark brown, 
appressed, 2 to 6 cells in length, with a pointed terminal cell and a spur at the base; 
rootlets densely villous. Stipes wingless, dark brown, dull, cylindrical, smooth or 
very slightly scabrous, terete, 3 to 11 cm. long, 0-2 to 0-8 mm. in diam., sparsely 
clothed towards the apex and base with similar hairs to those on the rhizome. Main 
rhachis dark brown, fiexuose, narrowly winged throughout, glabrous or sparsely 
clothed with a few, weak, appressed, dark brown, unbranched hairs about 4 to 6 


201 . Hymenophyllaceae. ( Tindale ) 


31 


cells in length. Fronds 8 to 35 cm. long including the stipes, pendulous in epiphytic 
plants. Lamina 3-pinnate-pinnatifid, submembranous to herbaceous, green when 
fresh, brown when dried, rather rigid, deltoid or lanceolate (3: 1) to very broadly 
ovate (6: 5), usually 5-5 to 12 cm. long, 2-5 to 7 cm. broad, rarely up to 15 cm. 
broad, the apex usually deflexed, acute or acuminate, the internal cell-walls smooth 
or crenately thickened. Primary pinnae with petiolules which are usually winged 
on the upper side, often upcurved, alternate, mostly 1 -2 to 6 cm. long, 1 -5 to 3 cm. 
broad, deltoid or ovate-lanceolate, the apex acute or obtuse. Secondary pinnae 
with winged petiolules, rhomboidal, alternate, again pinnatifid or 2-pinnatifid, the 
apex obtuse. Secondary rhachises winged throughout. Ultimate segments plane, 
cultrate (6: 1) to narrowly oblong (3: 1), the apex obtuse, the margin spinulose- 
dentate. Veins free, dichotomous, not reaching the apex of the ultimate segments, 
glabrous or with an occasional, red-brown, unbranched hair. Sori indusiate, 
marginal, numerous, mainly towards the apex of the frond, terminal on the short 
ultimate segments, immersed at the base. Involucre 0-8 to 1 mm. long, 0-5 to 1 mm. 
broad, orbicular to oval, entire, cuneate at the base, 2-valved almost to the base. 
Receptacle included or slightly exserted, conico-cylindrical. Sporangia turbinate, 
with an annulus consisting of 23 to 25 thick-walled cells and about 7 to 9 thin-walled 
cells. Spores globose-tetrahedral, trilete. minutely papillate, equat. diam. 45 to 
56-25[x X 41-25 to 52-5^. 

Distribution: Australia (south-eastern Queensland and the lower Central Coast of New 
South Wales) and New Zealand (including Stewart and Chatham Islands). 

Lectotype: “ No. 301.466, labelled Trichomanes bivalve , G. Forster’s Herbarium” with 
” New Zealand” added in pencil (BM), examined. This is a good match for D. A. Goy & 
L. S. Smith No. 72 which was collected at Roberts’ Plateau, Lamington National Park, south¬ 
eastern Queensland. I have also examined a fertile specimen of Hymenophyllum bivalve collected 
by Forster but without a specific locality (UPS) and a sterile fragment collected by Forster in 
the Society Islands (Herbar. Filic. Luerss. No. 10923), Kaulfuss No. 2601 (P). 

Queensland: Roberts' Plateau, Lamington National Park, very common as an epiphyte 
on tree trunks but also as a terrestrial plant among moss, Goy & Smith No. 72, 1.1938 (NSW 
PI286; BRI); ditto, on a tree trunk, Shirley (BRI); Moreton District. Lamington National 
Park. alt. about 3,000 ft., common on mossy fallen tree trunks in rainforest, Goy & Smith 
12.1942 (BRI); ditto, near Picnic Rock, Smith 12.1943 (BRI); Lamington National Park, 
Elbana Falls track, alt. ca. 2,800 ft., growing on the bases of tree trunks in rainforest, Smith & 
Tindale 8.1956 (NSW P7454); Lamington National Park, epiphytic on Nothofagus moorei 
between Bethongabel and Mt. Wanungra, Smith No. 10911, 3.1960 (BRI; NSW P8144). 

New South Wales: Minnamurra Falls, 3 miles W of Jamberoo, 1,200 ft. alt., on the 
face of a boulder, in rainforest, in the mist region, Judd 5.1954 (NSW P6786); above Minnamurra 
Falls, on the road to Robertson, alt. 2,200 ft., in cleft of rock, in a rainforest ravine, above the 
stream, Judd 5.1955 (NSW P7164); Cambewarra, Bauerlen (BRI); Broghcr’s Creek, near 
Broughton Creek (on the Shoalhavcn River), Bauerlen 7.1884 (NSW PI906). 

Absolute Synonyms: Trichomanes bivalve Forst. f.. Prodr. (1786) 84. Sphaerocionium 
bivalve (Forst. f.) Pr., Hymen. (1843) 33. Meringium bivalve (Forst. f.) Copcl. in Phil. Journ. 
Sci. LXVlI (1938) 44; Crookes & Dobbie, New Zealand Ferns, ed, 5 (1952) 58, 104. 

Equivalent Synonyms: Hymenophyllum spathulatum Colenso in Tasm. Journ. II (1844) 
184. Holotype: on living trees, shores of Waikare Lake, W. Colenso 12.1841 (not examined). 
Hymenophyllum pyriforme v.d. Bosch in Ned. Kruid. Arch. V, 3 (1863) 173, Holotype: New 
Zealand, d’Urville (H. Franq.), (B), not examined. 

Hymenophyllum bivalve is a rare species in New South Wales where it has been recorded 
from mountain rainforests on the lower Central Coast. Until it was collected by Judd at the 
Minnamurra Falls Reserve, 3 miles west of Jamberoo, N.S.W., in 1954, no specimens had been 
recorded from this State since Bauerlen’s collections were made seventy years previously. 

This fern is found at elevations of over 1,000 ft., usually in subtropical rainforests but also 
extending into the Nothofagus moorei rainforests especially in south-eastern Queensland. This 
G 99777—3 






32 


201 . Hymenophyllaceae. ( Tindate ) 


species often occurs in mist regions particularly on boulders in ravines but otherwise it is found 
as an epiphyte on living trees or old logs or less frequently it is terrestrial. 

The material of H. bivalve from New Zealand is very similar to that from eastern Australia. 
In Trans. Roy. Soc. N.Z. LXXXV (1958) 213, Brownlie recorded a chromosome count of n 22 
for a specimen collected at North Westland, New Zealand (Brownlie No. 137). 


III. THE GROUP OF TRICHOMANES 

The following 7 genera which occur in south-eastern Australia, are recognized 
as belonging to this group:— Microgonium, Crepidomanes, Gonocormus, Polyph/ebium, 
Macroglena, Cephalomanes and Selenodesmium. They are construed here in the 
sense of Copeland in Genera Filicum (1947). All these genera are characterized 
by tubular or obconic involucres mostly with a truncate or trumpet-shaped mouth, 
although in some species of Crepidomanes , e.g. C. bipunctatum (Poir.) Copel., it is 
distinctly 2-lipped. 

The Trichomanes-gro up of species is generally believed to have been reduced 
from the HymenophyHum-group due to their smaller size, as well as to the absence 
of true roots in some of the minute species and sometimes the xylem absent or 
reduced to one tracheid. The false veins in Microgonium and Crepidomanes are 
also thought to have been reduced from true veins. The sporangia in the 
Hymenophyl/um-group are large and usually almost spherical, whereas those of 
the Trichomanes-gxoup are minute, compressed bodies. 

According to Bower (The Ferns II (1926) 244) the prothallus of the 
Tric/tomanes-group of species is filamentous, branched, resembling a green Alga, 
with the archegonia produced at the end of short branches called archegoniophores. 
whereas in the HymenophyHum-group the prothallus is flattened, strap-like, never 
cordate, with antheridia and archegonia borne together on a lateral lobe. 


5. MICROGONIUM Presl 

Microgonium Presl. Hymen. (1843) 19, pi. 6; van den Bosch, Hymen. Javan. 
(1861) 5; Copel. in Phil. Journ. Sci. LXVII (1938) 61; Copel., Gen. Fil. (1947) 39. 

Hemipltlebium Presl sect. Microgonium Prantl, Hymen. (1875) 48. 

Copeland in Phil. Journ. Sci. LI (1933) 196, treated Hemiphlebium Presl as a group of 
Trichomanes L. In Ferns of Malaya (1954) 91, Holttum regarded Microgonium as a subgenus 
of Trichomanes. 

Tiny, epiphytic ferns. Rhizome creeping, filamentous, usually without 
roots, densely clothed with hairs. Fronds usually simple or sometimes lobed, often 
orbicular, suborbicular or obovate-cuneate, the surface glabrous, the vernation 
straight. Veins pinnate or flabellate, sometimes with an intramarginal strand. 
False vein/els (single rows of darker cells which are usually narrower than the ordinary 
cells of the lamina) always present. Involucre elongated, free or immersed, narrowly 
tubular to trumpet-shaped, the mouth dilated and entire or slightly 2-lipped. 
Receptacle usually exserted. Sporangia subsessile, compressed. Spores tetrahedral 
to globose-tetrahedral, often granulate. 

Type of Genus: Microgonium cuspidatum (Willd.) Presl (Basionym: 
Trichomanes cuspidatum Willd.) of the East African Islands. 


201 . Hymenophyltaceae. (Tindale) 


33 


This is a genus of about 12 species ranging from Africa to Tahiti, Formosa 
and Australia with 1 species in Central America. According to Copeland in Genera 
Filicum (1947) 40, the species of Microgonium might be derived from Crepidomanes 
by extreme reduction of the fronds which are mostly simple in Microgonium but 
pinnately compound in the other genus. The lamina in both genera is characterized 
by false veinlets or striae. Some species of Microtrichomanes , e.g. M. vitiense 
(Bak.) Copel., are closely related to Microgonium but lack the false veinlets. The 
plants of Microgonium spp. as a whole are more reduced in size than those of any 
other genus formerly included in Trichomcmes sens. lat. 

There are 4 species of Microgonium in the rainforests of tropical north-eastern 
Queensland but only one of them, viz. M. tahitense , is believed to extend to the 
subtropical North Coast of New South Wales. However M. bimarginatum v.d. B. 
occurs as far south as Eudlo which is about 50 miles north of Brisbane, Queensland. 

A chromosome count of n = 34 was made on Malayan material of 
Microgonium motleyi van den Bosch by Manton & Sledge in Phil. Trans. Roy. Soc. 
Lond., Ser. B, Biol. Sci. No. 654, CCXXXVIll (1954) 136. This tiny epiphytic- 
species extends to the rainforests of north-eastern Queensland. 1 have examined 
the holotype of Trichomanes sayeri F. Muell. et Bak. which was collected by Sayer 
at Trinity Bay, north-eastern Queensland (BR1 32774), and consider it to be identical 
with M. motleyi. 


*• Fronds sessile and peltate. Submarginal vein in the lamina absent. M. tahitense 1. 

1* Fronds stipitate and non-peltate. Submarginal vein in the lamina present. 

. M. bimarginatum 2. 


1. Microgonium tahitense ( Nadeaud ) Tindale in Contrib. N.S.W. Nat. Herb. 
FI. Ser. 201 (1963) 4. 

Habitat in tropical and subtropical rainforests on tree-trunks or on dripping 
rocks. Rhizome long-creeping, wiry, about 0-2 mm. in diam., densely tomentose 
with almost black, unbranched hairs up to 4 cells in length. Fronds sessile, 
overlapping, almost glabrous above, clothed fairly densely with dark red to black, 
unbranched hairs along the abaxiai surface of the veins, peltate, attached to the 
rhizome near the centre or towards the base of the lamina, appressed to the bark or 
surface upon which the plant is growing, 0 4 to 2-5 cm. in diam., oblate (4: 3) to 
orbicular (6: 6), thin, membranous, the surface wavy with concentric undulations, 
the margin not thickened, entire or often erose, easily torn. Veins numerous, 
radiating from the point of attachment to the rhizome, forked (sometimes several 
times), ending close to the margin. False veins present between the true veins 
especially towards the periphery of the lamina. Sori few, 1 to 3 on each frond, 
scattered, partly embedded in the lamina and partly exserted beyond the margin. 
Involucre tubular, elongated, coriaceous, 2 to 3 mm. long, the tube 0-4 to 0-5 mm. 
broad, the base cuneate, the rim much dilated, spreading, entire, slightly 2-lipped, 
about 0-5 to 1 -0 mm. in diam. Receptacle slightly exserted or not exserted. 
Sporangia subsessile, compressed-turbinate, with an oblique annulus composed of 
17 to 23 indurated cells and about 4 to 5 thin-walled cells. Spores tetrahedral to 
globose-tetrahedral, trilete, green, the exine granulate, equat. diam. 30 to 39-38[x 
X polar diam. 24-38 to 31 -88p.. (No. 201, PI. V, f. 3.) 

Distribution: Samoa, Tahiti, Fiji, Rarotonga, New Caledonia, New Hebrides, New 
Guinea, Amboina, Java and Australia (north-eastern Queensland probably extending to the 
North Coast of New South Wales). 





34 


201 . Hyntenophyllaceae. (Tindale) 


Holotype: Vallee de Papenoo, Tahiti, Nadeaud. 


Queensland: Palm Camp, Bellenden Ker, 
Creek, North Cairns, Bailey (NSW P2I36, P3754; 
(K); Range, Trinity Bay, on trees (BRI). 


Bailey 1889 (BRI, part of 30483); Freshwater 
BRI); Trinity Bay, Bailey ex Herb. Mueller 


Absolute Synonym: Trichomanes tahitense Nadeaud, Enum. des Plantes Indigenes de 
Pile de Tahiti (1873) 18. 


Equivalent Synonyms: Trichomanes pellaiuni Bak. in Journ. Linn. Soc. IX (1866) 336. 
i 8 f C non T. peltatum Poir. (1808), Holotype: Samoa, Powell No. 125.7-1864-(K), examined. 
Microgonium omphalodes Vieill. ex Fourn. in Ann. Sci. Nat., Scr. V, XVIII (1873) 25 i (published 
in svnonynty for Trichomanes peltatum Bak. and therefore illegitimate); Copel. in I nil. Journ. 
Sci LXVII (1938) 63, Holotype: Samoa, Powell No. 125, 7.1864 (K), examined. Trichomanes 
'nannosum Cesati in Rend. Ac. Napoli XVI (1877) 24. 28, Holotype: Terra de. Papuas, M 
Arfak a Putat, Beccari 10.1872 (RO), examined. Hemiphlebium peltatum (Bak.) Luerss. in Bot. 
Centralbl. XI (1882) 28, based on T. peltatum Bak. Trichomanes omphalodes C. Chr., 

Fil. (1906) 646; Copel. in Phil. Journ. Sci. LI (1933) 203, pi. 31,.f. 1-6. 


Ind. 


Both Trichomanes tahitense Nadeaud and Microgomum omphalodes were published in 
1873 but Fournier mentioned T. tahitense as perhaps being synonymous with M. omphalodes , 
so that Nadeaud's species must have been published first. 


The specimen at Kew collected by Bailey at Trinity Bay, Queensland, agrees perfectly 
with Fournier's specimen of Microgonium omphalodes obtained at Wagap, New Caledonia. I 
have not examined the other New Caledonian specimen of M. omphalodes cited by Fournier, 
viz. Neoua, Depl. (Deplanche) No. 175. He also cited Samoa, Powell No. 128 (l.e. the holotype . 
of Trichomanes peltatum Bak.) and Viti and Pelew, Luerssen. 


The material of Trichomanes pannosum Cesati which was sent on loan by courtesy of the 
Botanical Institute, Rome, Italy, was collected in the Arfak Mountains. New Guinea, and is 
typical M. tahitense not Microgonium motleyi v.d.B. as Copeland suggested m Phil. Journ. Sci. 
LI (1933) 202. 

Although M. tahitense is widely distributed through Malaysia, the Pacific Islands and to 
Australia, this species shows remarkably little variation over its range. According to Watts 
on page 3 of an unpublished manuscript in the National Herbarium of New South Wales, Sydney, 
there was a record of this fern from the northern rivers of New South Wales. This species, 
as Trichomanes omphalodes , is listed for New South Wales by Maiden & Betche in A Census 
N.S.W. Plants (1916) I, but I have not seen any material of this fern from the latter State, the 
only Australian specimens which I have examined, were collected in north-eastern Queensland. 
However, this species could be easily overlooked since it grows on the bark ot trees and on rocks 
with the rounded, sessile, peltate fronds lying close to the surface of the substratum. 


2. Microgonium bimarginatum van den Bosch, Hymen. Javan. (1861) 7, 
Copeland in Phil. Journ. Sci. LXVII (1938) 61. 

Habitat in tropical and subtropical rainforests, often in gorges, growing on 
trees or damp rocks. Rhizome long-creeping, branched, 0-1 to 0-2 mm. in diam. 
(not including the lomentum), densely clothed with dark red, often glandular-tipped, 
unbranched hairs. Stipes comparatively distant, 0-5 to 0*9 mm. long, about 0-1 mm. 
in diam., dark brown or black, clothed with dark red, often glandular-tipped, 
unbranched hairs for most of their length. Fronds 0-4 to 1 -5 cm. long including 
the stipes. Lamina 0-6 to 1*3 cm. long, 01 to 0*8 cm. broad, thin, membranous, 
very variable in shape, ranging from obovate, suborbicular, elliptical, oblong to 
urceolate, the apex broadly rounded, truncate, bifid or lobed, the base rounded oi 
cuneate, the margin glabrous, entire, sinuous, convoluted or irregularly lobed, 
with a single row of hyaline cells between the submarginal strand and the margin. 
Costa prominent, zig-zagged, clothed with a few, unbranched hairs which are usually 
2 to 3 cells long. Lateral true veins prominent, arising front the costa and usually 


201 . Hymenophyllaceae. ( Tindale ) 


35 


terminating in the prominent submarginal strand. False veins numerous, free, 
occurring between the true veins. Sori few, 1 to 3 on each lamina, terminal on 
the apices of simple fronds or embedded in the apical lobes of more complex fronds. 
Involucre tubular, coriaceous, 1-5 to 2-5 mm. long, the tube 0-3 to 0-4 mm. broad, 
the rim much dilated and 1 to 1*2 mm. in diam. Receptacle long-exserted. 
Sporangia subsessile, compressed, the oblique annulus composed of 18 to 20 
indurated cells and about 4 thin-walled cells. Spores globose-tetrahedral or 
tetrahedral, greenish, the exine evenly granulate, equat. diam. 22-5 to 31-88p. X polar 
diam. 18-75 to 28-13g. (No. 201, PI. V, f. 2.) 

Distribution: Ceylon, Malaya, New Guinea, Fiji, Samoa and Australia (north-eastern 
to south-eastern Queensland). 

Syntypes: 1) Ins. Ceylon, Thwaites No. 2986 (GH), an illustration of this syntype 
examined (see Phil. Journ. Sci. LI (1933) pi. 33, f. 1). 2) Ins. Fidchi (Fiji), Wilkes (not examined). 

Queensland: Mossman Intake, alt. ca. 100 ft., Smith No. 3978A, 9.1948 (NSW P8276); 
Street’s Gully, Kuranda, Watts 7.1913 (NSW P8277); Cairns district. Watts, 7.1913 (NSW 
P8342); near Lake Barrine, Atherton Tableland, Bailey 6.1899 (NSW P8279; BRI 32790); 
Harvey’s Creek, 300 ft. alt., on damp rocks, in heavy forest in creek gorge, Messmer 8.1954 
(NSW P6898); Frenchman’s Creek, Watts 7.1913 (NSW P8278; BRI 32781); Herberton, 
Waller 1908 (NSW P6899); headwaters of Tully River, in rainforest on trees, Messmer 9.1954 
(NSW P6948); Mt. Spec, in rainforest, Vessey 4.1962 (NSW P8361); Eumundi, Bailey & 
Simmonds 5.1892 (NSW P8280; BRI 32786); Eudlo Creek, Bailey 11.1891 (BRI 32785); 
Eudlo, Simmonds (BRI 32784). 

Absolute Synonym: Trichomanes bimarginatum van den Bosch in Ned. Kruid. Arch. V 
(1861) 143; Domin in Bibl. Bot. LXXXV (1913) 11, t. 3, f. 2; Copeland in Phil. Journ. Sci. 
LI (1933) 208, pi. 33, f. 1-4; Holttum, Ferns of Malaya (1954) 92. 

Equivalent Synonym: Trichomanes yandinense F. M. Bail, in Proc. Linn. Soc. N.S.W. V 
(1881) 30; F. M. Bail., Syn. Queensl. FI. (1883) 686, Holotype: Maroochie, Queensland, 
F. M. Bailey 7.1879 (BRI 30442), examined. 

A specimen collected in Java, Batavia, G. Paniisan, O.v. Buitenzorg, ca. 700 m., by 
Bakhuisen van den Brink No. 6197, 12.1933 (BRI 32791) agrees well with some of the Queensland 
specimens with unlobed fronds. Trichomanes yandinense represents a form in which there 
are usually several lobes towards the apex of each lamina. 

The most distinctive feature of Microgonium bimarginatum is the more or less continuous, 
submarginal false vein which is not present in either M. tahitense or M. motleyi. The shape of 
the fronds varies greatly even on the same rhizome, suborbicular and urceolate laminas occurring 
quite close to each other. 

This species has been recorded as far south as Eudlo which is about 50 miles north of 
Brisbane, Queensland. This tiny epiphyte is found in dense rainforest especially in north-eastern 
Queensland. 

M. bimarginatum is very closely related to Trichomanes baileyanum W. W. Watts which 
was published in Proc. Linn. Soc. N.S.W., XXXIX (1914), 1915, 758. The holotype of the 
latter species, which was collected at Josephine Creek, Bartle Frere, north-eastern Queensland, 
by Watts in July, 1913 (NSW PI 766), is located in the National Herbarium, Sydney. The 
fronds of this type are more regularly pinnatifid than in M. bimarginatum but they have the 
characteristic submarginal vein. BRI Nos. 32787-9 and 32792 closely approximate to the 
holotype of T. baileyanum. The latter species is smaller than but may be a form of Microgonium 
mindorense (Christ) Copel. which is found in the Philippines and the Admiralty Islands. 


6. CREPIDOMANES Presl 


Crepidomanes Presl, Epim. Bot. (1851) 258; Copel. in Phil. Journ. Sci. LXVII 
(1938) 58; Copel., Gen. Fil. (1947) 39. 

G 99777—4 


36 


201 . Hymenophyllaceae. ( Tindale ) 


Taschneria Presl, Epim. Bot. (1851) 258; Copel. in Phil. Journ. Sci. LI (1933) 174, as 
a group of Trichomanes L. 

Didyrnoglossum Prantl, Hymen. (1875) 45, non Desv. 

In the Ferns of Malaya (1954) 99, Holttum treated Crepidomanes as a subgenus of 
Trichomanes. 

Small, epiphytic, rupestral or rarely terrestrial ferns. Rhizome filiform, 
widely creeping, clothed with hairs, mostly without roots. Axes usually winged. 
Fronds pinnately decompound, pinnate or sometimes digitate, the surface glabrous 
or almost so. Segments narrow, mostly with irregularly scattered, sclerenchymatous 
false veins or striae and/or with a submarginal vein, the margin entire and glabrous. 
Veins free, often clothed with appressed 2- or 3-celled hairs. Sori axillary or placed 
behind one another. Involucre campanulate, trumpet-shaped or obconic, often 
broadly winged, bilabiate or in C. christii with an entire, dilated mouth. Receptacle 
exserted or sometimes not exserted. Sporangia subsessile, compressed. Spores 
tetrahedral or globose-tetrahedral, trilete, greenish. Gametophyte filamentous, 
with the archegonia borne on archegoniophores, the neck of the archegonium 
straight and usually 4 cells long, the antheridia with a basal disk-like or wedge-shaped 
cell (occasionally with 2 basal cells), dehiscing by the discharging of the opercular 
cell. 

Type of Genus: Crepidomanes brevipes (Presl) Copel., (syn. Didyrnoglossum 
brevipes Presl), the Philippines, Guam and Borneo. 

There are 12 or more species most of them very variable and dwarfing is 
common. This genus ranges from the East African Islands to Japan, China, 
Queensland and Polynesia. One uncommon species Crepidomanes walleri (Watts) 
Tindale extends from the Atherton Tableland, north-eastern Queensland to 
Springbrook in south-eastern Queensland. There are at least 4 other species ot 
Crepidomanes in the tropical rainforests of north-eastern Queensland. They are 
* Crepidomanes barnardianum (F. M. Bail.) Tindale, C. venulosum (Ros.) Copel., 
C. majorae (Watts) N. A. Wakef. (only known from Ravenshoe, Queensland) 
and C. hipunctatum (Poir.) Copel. However, the tropical Australian material of 
this genus is badly in need of revision. 

Crepidomanes is allied to Microgonium which is also characterized by false 
veins or sclerenchymatous striae in the lamina. Copeland in Phil. Journ. Sci. 
LXVII (1938) 59, suggests that Crepidomanes is the parent genus of Microgonium 
which is composed of reduced species. 

Stokey made a detailed study of the gametophyte of a member of this genus, 
viz. Crepidomanes bilabiatum (Nees et Bl.) Copel., as Trichomanes bilabiatum Nees 
et Bl., in “ Reproductive Structures of the Gametophytes of Hymenophyllum and 
Trichomanes ” in Bot. Gaz. CIX (1948) 363-380. 

Chromosome counts have been made for several species of Crepidomanes 
including a count of n = ca. 36 for Malayan material of C. bipunctatum by Manton 
& Sledge in Phil. Trans. Roy. Soc. London, Ser. B, Biol. Sci. No. 654, 
Vol. CCXXXVIII (1954) 136. Manton & Sledge (l.c. 136) also recorded a count 
of n = 36 for material of C. bilabiatum collected at Hakgola, Ceylon. In both 
species the specimens were stated to be tetraploid. Chromosome counts on 2 
Indian species of this genus were recorded by Mehra & Gurdip Singh in “ Cytology 
of Hymenophyllaceae ” in Journ. Genetics LV (1957) 391. They were as follows:— 


Footnote: Crepidomanes barnardianum (F. M. Bail.) Tindale comb. nov. (Basionym : 
Trichomanes barnardianum F. M. Bail., Queensl. Flora Suppl. Ill (1890) 89). 



37 


201 . Hymenophyllaceae. ( Tindale ) 

Ci epidomanes plicatum (v.d. Bosch) Ching n = 36; C. insigne (v.d. Bosch) Fu var. 

ul A'J nSl A- ne ^ V \ T an< ^ C. insigne var. y 2n = 72. According 
to JVLehra and Gurdip Singh C. insigne is a species complex which requires taxonomic 
research. 


/1 £ r , epi n tlomanes walleri ( Watts ) Tindale in Contrib. N.S.W. Nat. Herb. FI. Ser.. 
2U1 (1963) y. 

Habitat in tropical and subtropical rainforests on mossy rocks. Rhizome 
long-creeping, shortly branched, 0.1 to 0.3 mm. in diam. (not including the 
tomentum), densely clothed with hairs which are line, dark brown, unbranched, 
septate, mostly 3 to 6 cells in length and sometimes with a small, round, 
glandular-tipped apex. Stipes not winged or very shortly winged towards the apex, 
to .} cm - |p n 8» clothed with an occasional, clavate, fawn, 2-celled hair towards 
the middle and apex but towards the base with similar hairs to those on the rhizome. 
Main rhachis winged by the decurrent bases of the primary pinnae. Fronds 1 to 
4 cm. in length including the stipes. Lamina 1-pinnate-pinnatifid to almost 2-pinnate, 
sometimes 1-pinnate, 0-5 to 3 cm. long, 0-8 to 1-5 cm. broad, thin, membranous, 
irregularly narrowly elliptical (3: 1) to broadly elliptical (3: 2) or very broadly ovate 
(6: 6) to lanceolate (3 : 1), the apex acute or rounded, the surface glabrous except 
the veins. Primary pinnae either simple and linear to cultrate or ± flabellate and 
irregularly 2- to 5-lobed, 5 to 10 mm. long, 0-5 to 6 mm. broad. Secondary segments 
not petiolulate, linear (10: 1) to broadly oblong (3:2), the apex obtuse, truncate 
or emarginate, the margin entire or slightly undulate. Veins free, each laminal 
segment with a single vein which does not reach the apex, lacking a continuous 
vein near the margin, clothed especially on the adaxial surface with hairs which 
are appressed, fawn or dark brown, clavate, the large apical cell often darker. 
False veins numerous, short, flexuose, very divergent from the costa. Sori few, 
1 to 8, embedded in the apices of short, basal, acroscopic segments of the primary 
pinnae. Involucre obconical with 2 large acutely or obtusely triangular lips, green, 
with false veinlets towards the apex and base. Receptacle not exserted or shortly 
exserted. Sporangia subsessile, compressed, with an annulus composed of 15 to 
19 indurated cells and 3 to 4 thin-walled cells. Spores tetrahedral, trilete, greenish, 
minutely pitted, equat. diam. 37-5 to 45^ X polar diam. 31-88 to 37-5 (a. 

Distribution: Australia (North-eastern Queensland to the McPherson Range in south¬ 
eastern Queensland). 


Holotype: Herberton district, Waller 1908 (NSW P1311). 

Queensland: Evelyn Scrub, Herberton district. Waller No. A3, 11. 1908 (NSW P834r); 
Johnstone River, Kefford (BRI 28921; NSW PI312); Wide Bay district, Mt. Cooroy, on mossy 
rocks in rainforest, uncommon, Wakefield 3.1943 (BRI 28923); Eudlo, Simmonds (BRI 28919; 
NSW P8343); Nerang Creek, Springbrook, Schneider 7.1884 (BRI 28922). 

Absolute Synonym: Trichomanes walleri Watts in Proc. Linn. Soc. N.S.W. XXXIX, 
1914 (1915) 761, pi. lxxxvi, f. 3 a-d. 


This species is closely allied to Crepidomanes majorae (Watts) N. A. Wakef. which has 
been collected only at the type locality, viz. Ravenshoe, North Queensland. However, the 
mvolucre in C. majorae is tubular with a scarcely dilated, non-bilabiate apex whereas it is obconical 
with 2 lips in C. walleri being very similar to that of C. bipunctatum. The intramarginal vein 
is missing in the lamina of both C. majorae and C. walleri whereas it is very prominent in C. 
bipunctatum. The latter species has much larger fronds, i.e. up to 7-5 cm. long including the 
stipes, than in C. majorae or C. walleri where they range from 1 to 4 cm. long. The false veinlets 
in C. walleri are much more divergent from the costa than in C. majorae where they are almost 
parallel to the midrib. 


38 


201 . Hymenophyllaceae. ( Tindale ) 


7. GONOCORMUS van den Bosch 

Gonocormus van den Bosch, Hymen. Javan. (1861) 7; van den Bosch, Eerste 
Bijdrage (1861) 321; Copel. in Phil. Journ. Sci. LXVII (1938) 56-7; Copel., Gen 
Fil. (1947) 38. 

Copeland in Phil. Journ. Sci. LI (1933) 143, referred to Gonocormus as a group of 
Trichomanes L. In Ferns of Malaya (1954) 95, Holttum treated Gonocormus as a subgenus of 
Trichomanes following Christensen in Ind. Fil. (1906) 634. 

Small, epiphytic or rupestral ferns forming large mat-like patches. Rhizome 
filamentous, much branched, proliferous, densely clothed with unbranched hairs. 
Stipes and rhachises filamentous, much branched, dark, wiry, often bearing proliferous 
buds. Fronds small, flabellate or circular, dark green, glabrous or almost so, often 
pinnate. Segments narrow, with flabellate venation, false veinlets absent, the 
margin unthickened and incised, the apices usually emarginate or bifid, the cell-walls 
thin and unpitted. Involucre sunken in the apices of the segments, tubular, with an 
entire, dilated mouth. Receptacle extruded in the older stages. Sporangia 
compressed, the annulus almost transverse, composed of 15 to 19 indurated cells 
and about 3 or 4 thin-walled cells. 

Type of Genus: Gonocormus minutus (Bl.) van den Bosch (Basionym: 
Trichomanes minutum BL), Africa to Java and Polynesia extending to Japan and 
Australia. 

This is a small and rather distinctive genus of several poorly defined species, 
2 being found in Australia. Gonocormus saxifragoides (Pr.) van den Bosch extends 
from the Atherton Tableland of north-eastern Queensland to the North Coast of 
New South Wales, whereas G. minutus occurs in the tropical rainforests of north¬ 
eastern Queensland. 

In this genus the axes of the frond and the rhizome are poorly differentiated 
and with little distinction of function, as has been indicated by E. B. Copeland in 
Genera Filicum (1947) 38. Buds on the fronds may develop into new fronds or 
even into a rhizome. In Ferns of Malaya (1954) 95, Holttum has suggested that 
the members of the genus (or subgenus in his opinion) are possibly reduced relatives 
of Vandenboschia. 


Gonocormus saxifragoides ( Presl ) van den Bosch, Hymen. Jav. (1861) 9. 

Habitat in tropical or subtropical rainforests either epiphytic on tree-trunks 
or more frequently growing in patches on wet rocks. Rhizome long-creeping, much 
branched, about 0-1 mm. in diam. (not including the tomentum), densely clothed 
with hairs which are simple, brown, often glandular-tipped and usually 3 to 6 cells 
in length. Stipes dark, wiry, 3 to 20 mm. long, about 0-1 mm. broad, densely 
tomentose for one-third to two-thirds of their length with similar hairs to those 
on the rhizome. Fronds 0-5 to 2-5 cm. long including the stipes, mostly simple, 
rarely proliferous. Lamina flabelliform, reniform, almost circular or ovate-cuneate, 
dark green, fluted, channelled in the dried condition, 3 to 10 mm. long, 6 to 11 mm. 
broad, often deeply incised between the groups of veins but more shallowly between 
the ultimate lobes, the cell-walls wavy. Segments more or less oblong, 1 to 7 mm. 
long, 0-3 to 0-7 mm. broad, the medial segments usually longer than the lateral 
ones, the apices obtuse and emarginate or bifid. Veins flabellate, without false 
veinlets. Sori immersed in the apices of the segments, 1 to 6 on each frond. 
Involucre 1*5 to 2 mm. long, 0-2 to 0-3 mm. broad at the base, narrowly urceolate. 


201 . Hymenophyllaceae. ( Tindale ) 


39 


the tube elongated, the mouth broadly dilated and entire, clothed towards the base 
with a few, dark red or brown, glandular-tipped hairs which are 1 to rarely 6 cells 
in length. Receptacle narrow, included or shortly extruded in the later stages. 
Sporangia compressed, the annulus almost transverse, composed of 17 to 19 indurated 
cells and about 3 or 4 thin-walled cells, without a definite stomium, dehiscing by 
a lateral slit. Spores globose-tetrahedral or tetrahedral, greenish, finely 
granulate, equat. diam. 30 to 56-25pt x polar diam. 28 to 54-38g. 


Distribution: Africa to Java and Malaya extending to Japan, Polynesia and Australia 
(Coast and Tablelands from north-eastern Queensland to the North Coast of New South Wales). 

Holotype: Luzon, the Philippines, Cuming No. 256 (K), examined. 

Queensland: Iron Range, 200 miles N of Cooktown, Flecker 4.1944 (NSW P8275); 
Street’s Gully, Kuranda, Watts 7.1913 (NSW P2202, P3779); Frenchman’s Creek, near Cairns, 
Watts 7.1913 (NSW P2204. P3791); Babinda Creek, 300 ft. alt., in rainforest on wet rocks, 
Messmer 8.1954 (NSW P6878); Babinda to Russell River, Watts 7.1913 (NSW P3781); Deeral, 
near Babinda, in dense rainforest on a rather steep slope, forming dense, deep green masses on 
rocks, about 350 ft. alt., Blake No. 14968, 7.1943 (BRI 28912; NSW P5271); between Cairns 
and Herberton, Wild 1891 (BRI 28900); Bartle Frere, Watts 7-8.1913 (NSW P3782; BRI 
28903); Herberton, Waller 1908 (NSW P2205); Eumundi, Bailey & Simmonds 11.1894 (BRI 
28907); Yandina, Bailey 3.1891 (BRI 28898); Yandina, Simmonds 3.1891 (BRI 28911; NSW 
P5279); Eudlo Scrubs, Bailey 11.1891 (BRI 28899); Ithaca Creek, near Brisbane, on rocks, 
Bailey (BRI 28906, 28910); Three Mile Scrub (Brisbane), Bailey (BRI 28918); Mt. Mistake 
(Lockyer Valley), Bailey (NSW P581; BRI 28908). 

New South Wales: Tweed River, Forsyth, probably about 1900 (NSW P2203); ditto, 
Dallachy 1.1892 (NSW P1861); Brunswick River, Watts 1899 (NSW P3780); head of Teven, 
Richmond River, on a rosewood tree. Watts 5.1899 (NSW P3785); Marshall Falls, Alstonville, 
Richmond River, Bauerlen 5.1891 (NSW P2200); Uralba, Richmond River, on trees, Watts 
1898 (NSW P3788); Three Mile Scrub, Richmond River, Watts 11.1902 (NSW P3789). 

Equivalent Synonym: Trichomanes saxifragoides Presl, Hymen. (1843) 39. 

Many authors referred to Gonocormus saxifragoides as Trichomanes parvulum J. Sm. 
non Poir. e.g. J. Sm. in Journ. Bot. Ill (1841) 417; Hook., Sp. Fil. I (1846) 118, pi. 39A; Benth., 
FI. Austr. VII (1878) 701; Bail., Lith. Ferns Queensl. (1892) 22, top figure; Moore & Betche, 
Handb. FI. N.S.W. (1893) 503; Domin in Bibl. Bot. LXXXV (1913) 11; Maiden & Betche, 
Census N.S.W. PI. (1916) 1; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114, and Copeland 
in Phil. Journ. Sci. LI (1933) 145-6. Later, in Phil. Journ. Sci. LXVII (1938) 37 and 56 Copeland 
agreed with Christensen that the true Trichomanes parvulum is a species now known as 
Hymenophyllum sibthorpioides (Bory) Mett. This problem was discussed by Christensen in 
Dansk Bot. Arkiv VII (1932) 3. 

According to Copeland in Fern Flora of the Philippines I (1958) 69, the small, almost 
non-proliferous form with circular fronds is equivalent to the original Trichomanes saxifragoides 
Presl and is produced in dry seasons, whereas the form with larger, more proliferous fronds 
is produced in wetter seasons. However, there appear to me to be 2 distinct species in Eastern 
Queensland. In Gonocormus saxifragoides which extends from tropical north-eastern Queensland 
to the subtropical North Coast of New South Wales as far south as the Richmond River, the 
fronds are seldom proliferous but are broader and rounder than in the other species which is 
confined in Australia to the rainforests of north-eastern Queensland. The latter species (i.e. 
Gonocormus minutus) is characterized by more finely divided, more delicate, mostly proliferous 
fronds in which there are very narrow wings on the axes and soriferous portions. I have not 
examined any material of G. minutus collected south of Babinda which is about 1,200 miles 
north of Brisbane, Queensland. 

Material of G. saxifragoides, which was very kindly collected by Dr. L. J. Webb and 
Mr. G. J. Tracy at The Boulders, Babinda, North Queensland, and cultivated in one of the hot 
houses at the Royal Botanic Gardens, Sydney, has not altered in the texture of its laminas after 
6 months nor has it produced any proliferous fronds. Plants of G. minutus collected near 
Mareeba, North Queensland, by Dr. H. S. McKee have also been cultivated at the Royal Botanic 
Gardens, Sydney. These fronds are much more delicate and translucent than in the material 
of G. saxifragoides collected at The Boulders by Webb and Tracy. 


40 


201 . Hymenophyllaceae. ( Tindale ) 


G. saxifragoides does not appear to have been collected in New South Wales since 1902; 
attempts to obtain this fern by myself and others in the rainforests of the North Coast have 
been unsuccessful. Mr. E. F. Constable carefully searched the Marshall Falls district, near 
Alstonville, and other localities in which this species was formerly recorded. 

The holotype of G. minulus was obtained on loan by courtesy of The Director, the 
Rijksherbarium, Leiden, for comparison with the Australian material of Gonocormus. This 
specimen which was collected in Java by Blume, has much thinner, narrower and more diffuse 
laminas than in G. saxifragoides. Even the simple fronds of G. minutus (e.g. NSW P6950 
collected at Harvey’s Creek, north of Bellenden-Kcr, North Queensland, by Messmer) have 
more delicate laminas than those of G. saxifragoides. 


8. POLYPHLEBIUM Copel. 

Polyphlebium Copel. in Phil. Journ. Sci. LXVII (1938) 55. 

PhlebiophyUum van den Bosch in Versl. Akad. Wet. Amsterdam XI (1861) 321, non 
Phlebophyllurn Nees (orthographic variants). 

In Phil. Journ. Sci. LI (1933) 138, Copeland treated Polyphlebium as a monotypic group 
of Trichomanes i.e. the group of PhlebiophyUum. 

A monotypic genus occurring in Australia (south-eastern Queensland, New 
South Wales, Victoria and Tasmania), New Zealand, the Chatham, Stewart and 
Kermadec Islands. 

Type of Genus: Polyphlebium venosum (R. Br.) Copel., based on Trichomanes 
venosum R. Br. 

Polyphlebium venosum (/?. Br.) Copel. in Phil. Journ. Sci. LXVII (1938) 55; 
Crookes & Dobbie, New Zealand Ferns, ed. 4 (1951) 112, 113 (photo). 

Habitat in temperate and subtropical rainforests forming densely matted 
patches usually on trees, logs or tree-ferns (especially Dicksonia spp.). Rhizome 
widely creeping, branched, very dark brown or brown, about 0-2 to 0-3 mm. in 
diam., very densely clothed with a tomentum of ferruginous hairs. Stipes not 
winged, 1 to 5-5 cm. long, filiform, approximate or distant on the rhizome, slightly 
flattened, 1-sulcate above, light brown or light red-brown when dried, green in 
fresh condition, dark brown at the base, wiry, dull, often minutely scabrous, slightly 
tomentose at the base, otherwise glabrous or with a few, short, unbranched, scattered 
hairs. Rhachis broadly winged towards the apex, wingless below, usually glabrous 
or very sparsely hirsute. Lamina pinnatifid or pinnate-pinnatifid, membranous, 
pellucid, pale green, very irregular in outline, elliptical (2:1) to rounded (6: 5), 
narrowly lanceolate (6: 1) to broadly ovate (6: 5), narrowly oblanceolate (6: 1) 
to broadly obovate (6: 5), 2-5 to 10-5 cm. long, rarely up to 15 cm. including the 
stipes, 0-5 to 5-5 cm. broad, rarely 6-5 cm. broad, not dimorphic, glabrous or with 
a few, scattered, unbranched hairs, the internal cells of the lamina with straight 
walls. Primary pinnae petiolulate in the lower segments, broadly sessile above, 
2 to 9 pairs, alternate throughout or the lower pairs opposite, falcate, linear (12: 1) 
to cultrate (6: 1), ovate (3: 2), rhomboidal or flabellate, usually 0-5 to 3-5 cm. long 
and 0-2 to T5 cm. broad, the lowest 1 or 2 pairs often reduced to 1 to 3 mm. in 
length; apex obtuse, truncate or emarginate; margin crenate or sinuate, glabrous, 
not specialized. Ultimate segments more or less oblong except the apical segment 
which is usually linear (12:1 or more); apex emarginate or truncate. Veins 
reticulate, very dark brown, prominent; midrib flexuose; lateral veinlets alternate, 
once to several times forked in each ultimate segment or lobule; false veinlets 
absent. Sori marginal, few, 2 to 14 per frond, occurring towards the base as well 


201 . Hymenophyllaceae. ( Tindale ) 


41 


as towards the apex of the lamina, usually solitary on each primary pinna, partly 
sunken in a short, upper, basal lobe, but in large fronds up to 4 sori occurring on 
the upper and lower margins; each sorus terminal on a minor veinlet. Involucre 
1-5 to 2 mm. long, about 0-3 to 0-5 mm. broad, tubular, green, the mouth broadly 
dilated, entire or very slightly 2-lipped. Receptacle slender, cylindrical, long- 
extruded, sometimes projecting 2-5 to 8 mm. beyond the rim. Sporangia 20 to 50 
per sorus, subsessile, overlapping on the receptacle, broader towards the apex than 
the base, compressed, the annulus large and oblique, composed of about 16 to 19 
indurated cells and ca. 3 thin-walled cells, with obliquely lateral dehiscence. Spores 
globose-tetrahedral to globose, trilete, green, the exine thin and subgranulate, the 
long laesurae with narrow, subgranular ridges, equat. diam. 22-5 to 31 -88 h- X polar 
diam. 22-5 to 31 -88ft. Gametophyte *: delicate and long-lived; the spore germination 
is a modification of the 4-celled type in which there is a delay or suppression of the 
formation of walls in 1 or 2 of the tips; after germination a rhizoid appears and 
several weeks later a filament cell develops; after 3 months another rhizoid or 
sometimes another filament develops; after 5 months the prothallus is a monoecious, 
uniseriate, branching filament of about 20 cells with 2 or 3 rhizoids; the archegonia 
(with 4 to 6 rows of neck cells and straight necks) are borne on the sessile or stalked 
archegoniophores, the small, numerous, stalked antheridia each with a simple wall 
and an operculum which is raised or shed; regeneration by gemmae or fragmentation; 
embryo with a small haustorial foot but without a primary root, the rhizoid-like 
hairs developing early; several embryos even in the same archegoniophore. (No. 
201, PI. IV.) 

Distribution: Australia (south-eastern Queensland, New South Wales, Victoria and 
Tasmania), New Zealand, the Chatham, Stewart and Kermadec Islands. 


Holotype: Derwent, Ins. Van Diemen, R. Brown No. 96, 1804 (BM), examined. Isotype 
(K). There is also another specimen at the Kew Herbarium labelled “ Van Diemen’s Island, 
R. Brown ”. 

Queensland: Roberts Plateau, Shirley (BRI); Roberts Plateau, Lamington National 
Park, densely covering the trunks of Cyathea leichhardtiana, Smith & Goy No. 69, 1.1938 (BRI; 
BM); Moreton District, Lamington National Park, in antarctic beech forest (Nothofagus, etc.), 
3,700-3,800 ft. alt., chiefly epiphytic on Dicksonia , Blake No. 12978, 5.1937 (BRI); ditto, alt. 
3,500 ft., Smith 12.1942 (BRI); National Park, McPherson Range, White 1-2.1917 (BRI); 
McPherson Range, alt. 3,500 ft., White 1.1919 (NSW P586). 

New South Wales: Clarence River, Wilcox 11.1875, ex Herb. Mueller (K); Happy 
Valley, Mt. Wilson, alt. 3,000 ft., basalt moist gully, growing on the trunk of Dicksonia, Constable 
12.1948 (NSW P5641); ditto. Watts 12.1914 (NSW PI 155); ditto, on trunks of Dicksonia 
antarctica, in rainforest, basalt, Tindale & Constable 3.1952 (NSW P2871); Waterfall Gully, 
Mt. Wilson, epiphytic on Dicksonia antarctica, at the edge of a creek, rainforest, Melville & 
Tindale 4.1953 (NSW P6405); Jenolan Caves, Blakely 6.1899 (NSW P2099); Katoomba, 
Forsyth 12.1899 (NSW P2102); Lcura Glen, on a tree-fern. Watts 4.1903 (NSW PI 156); Leura 
Falls. Watts 4.1903 (NSW PI 153); ditto, Stephenson 1.1890 (NSW P2107) and 12.1890 
(NSW P5645); Mittagong, Woolls (NSW P2103); Minnamurra Falls, 3 miles W of Jamberoo, 
1,200 ft. alt., in rainforest, on a boulder, in the mist region, Judd 5.1954 (NSW P6787); 
Cambewarra Mountain Road, Watts 5.1903 (NSW PI 154); Sugar Loaf Mountain, Boorman 

I. 1915 (NSW P2109); $ mile NE of Trig., Mt. Dromedary, Central Tilba, plentiful on tree-fern 
trunks in rainforest, Constable 9.1953 (NSW P6640). 

Victoria: Gippsland, F. Mueller, ex Herb. Hooker 1867 (K); Mt. Juliette, source of 
Yarra, ex Herb. Mueller 7.1877 (K); Fernshaw, Deane 6.1889 (NSW PI 157); Etta’s Glen, 
Blacks Spur, Healesville, Watts 12.1906 (NSWP1152); Dandenong Ranges, E of Melbourne, 
epiphytic mainly on tree-fern trunks, fronds pendulous, Smith No. M100, 11.1943 (BRI); 
Dandenong Ranges, Mueller 1897 (K); Gembrook, Staer 4.1911 (NSW P2100); Lome, Watts 

II. 1919 (NSW PI 150, P5324); Otway Ranges, Williamson 12.1903 (NSW P2104). 


* Adapted from Stone in Aust. Journ. Bot. VI (1958) 183-203. 



42 


201 . Hymenophyltaceae. ( Tindale ) 


Tasmania: North-East: Mt. Barrow, Co. Dorset, Rupp 1.1922 (NSW P2108); St. 
Patrick’s River, on fern-trees, Gunn No. 1515 , 4.1845 (NSW P2101). West Coast: Kelly’s 
Basin, Macquarie Harbour, Milligan No. 911, received 9.1868 (K). South-East: National 
Park, Coombe No. 1794, 12.1929 (HO H1274); Russell Falls, on Dicksonia trunk, Johnson 
1.1949 (NSW P5672); near Hobart, Le Tall, about 1896 (K); Table Mountain, Van Diemen’s 
Island, Caley 11.1804 (BM); Fern Tree Hill, Caley 11.1804 (BM). 

Absolute Synonym: Trichomanes venosum R. Br., Prodr. FI. N. Holl. (1810) 159; Hook. 
& Grev., Ic. Fil. I (1828) t. 78; Benth., FI. Austr. VII (1878) 702; F. M. Bail., Lith. Ferns 
Queens!. (1892) t. 23; Moore & Belche, Handb. FJ. N.S.W. (1893) 503; Rodway, Tasm. FI. 
(1903) 289; C. Chr„ Ind. Fil. (1906) 651; Domin. Bibl. Bot. LXXXV (1913) 14; Maiden & 
Betche, Census N.S.W. PI. (1916) 1; Ewart, FI. Viet. (1931) 31; Copel. in Phil. Journ. Sci. LI 
(1933) 138; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114. 

Equivalent Synonym: Trichomanes venustulum ( venustula ) Colenso in Trans. & Proc. 
N.Z. Inst. XII (1879) 1880, 366. Holotype by monotypy, on trunks of living trees, dense 
shady damp forests, west slopes of Ruahine Mountain Range, head of the River Manawatu, 
W. Colenso 1878-9 (not examined). Isotype: New Zealand, W. Colenso 9.1883, labelled 
T. venustula by Colenso (K), seen. This is a good match for NSW P2108 from Mt. Barrow, 
Tasmania. This form has more numerous secondary segments but does not appear to be 
worthy of specific rank. In some of the larger Australian specimens the sori are scattered on 
both edges of the pinnae. 

The monotypic genus of Polyphlebium is probably most closely related to the group of 
Vandenboschia pyxidifera (L.) Copel. The involucres are very similar but there is no branching 
of the veins in the undivided segments of the lamina in the latter group. 

In south-eastern Australia Polyphlebium venosum is a fairly common epiphytic fern which 
usually grows on the sheltered side of the trunks of Dicksonia antarctica but is sometimes found 
on the caudices ot Cyatheaaustralis, C. leichhardtiana and Dicksonia youngiae. According to 
Stone in Aust. Journ. Bot. \I (1958) 187, the gametophytes grow more readily among the root 
hairs of D. antarctica than on the caudices of other tree-ferns. P. venosum ranges from the 
subtropical rainforests and Nothofagus moorei forests of the McPherson Range, south-eastern 
Queensland, to the temperate rainforests of Victoria and Tasmania where Nothofagus 
cunninghamii is the dominant species. In eastern New South Wales P. venosum is mainly found 
m the subtropical rainforests especially in the Blue Mountains and on the South Coast. 
Frequently it grows in association with Hvmenophyllum flabellatunt which has a very similar 
range in Australia. 

There does not appear to be any difference between the specimens of P. venosum from 
New Zealand and those from Australia. In Trans. Roy. Soc. N.Z. LXXXII (1954) 665, Brownlie 
recorded a chromosome count of n = 36 for material of this species collected in New Zealand. 


9. MACROGLENA (Presl) Copel. 

Trichomanes L. sect. Macroglena Presl in Abh. Bohm. Ges. Wiss. V, 5 (1848) 
333. Macroglena (Presl) Copel. in Phil. Journ. Sci. LXVII (1938) 82, as a genus: 
Copel. in Gen. Fil. (1947) 44. 

Copeland in Phil. Journ. Sci. LI (1933) 258, treated Macroglena as a group of Trichomanes, 
whereas Holttum in Ferns of Malaya regarded it as a subgenus of Trichomanes. 

Small epiphytic or terrestrial ferns. Rhizome short, thick and erect or long- 
creeping, 1 to 10 mm. in diam., clothed with dark red or purplish-brown, bristle-like 
hairs. Stipes fascicled or scattered along the rhizome. Lamina pinnately 
decompound, very finely dissected. Ultimate segments very narrow, 1-veined, 
often stiff and bristle-like, otherwise soft, the cell-walls either thickened and coarsely 
pitted or thin and unpitted, the cells large and usually with brownish contents 
appressed to the walls. Involucre cupuliform to almost triangular, the mouth 
mostly truncate and entire, otherwise slightly expanded. Receptacle long, exserted. 
Sporangia compressed, subsessile. Spores globose-tetrahedral to globose. 


201 . Hymenophyllaceae. ( Tindale ) 


43 


Type of Genus: Macroglem meifolia (Bory) Copel. (syn. Trichomanes meifolia 
Bory), native to Madagascar, Reunion, Indonesia, Malaya to Samoa. 

This genus ranges from the East African Islands to Indonesia, Polynesia, 
Eastern Australia and New Zealand. Its principal features are the extreme reduction 
of the laminal segments, the coarsely pitted cell-walls of most species and the brownish 
laminal cells. Some species show marked affinities with Selenodesmium, others 
with Abrodictyum according to Copeland. 

In Eastern Australia there are 2 species of Macroglena, viz. M. caudata 
(Brack.) Copel., which ranges from the tropical rainforests of north-eastern 
Queensland to the temperate rainforests of Gippsland, Victoria, and M. meifolia 
which occurs in north-eastern Queensland. In the latter species the laminal segments 
are very fine and bristle-like, with only 1 or 2 rows of cells on each side of the veins. 
This tropical species is sometimes found totally submerged in running streams, 
e.g. Brass & White No. 236, collected on Thornton’s Peak, Queensland, at 4,000 ft. 
alt. (BRI 30371). 


Macroglena caudata {Brack.) Copel. in Phil. Journ. Sci. LXVII (1938) 84; 
Wakef., Ferns of Viet. & Tas. (1955) 4 f. 8, 7. 

Habitat epiphytic on the trunks of tree-ferns (e.g. Dicksonia spp., Cyathea 
australis and C. leichhardtiana) and trees in subtropical, tropical and temperate 
rainforests. Rhizome long-creeping, dark brown, flexible but not wiry, 1 to T5 mm. 
in diam., densely clothed with linear, spreading, bristle-like, dark red-brown hairs 
with narrow constrictions between the cells. Stipes wingless except for 2 almost 
imperceptible ridges which are continuous throughout the length of the stipes, 
T5 to 7 cm. long, 0-3 to 0-8 mm. broad near the middle, scattered along the rhizome, 
usually 1 to 4 cm. apart but crowded towards the apex of the rhizome (i.e. about 
1 mm. apart), slender, terete, slightly scabrous, glabrous except for a few dark 
red-brown, bristle-like hairs at the base. Main rhachis winged almost imperceptibly 
throughout its length. Lamina 2- to 3-pinnate-pinnatifid, membranous, pellucid, 
medium green or often brown when dried, very narrowly elliptical (6: 1) to narrowly 
elliptical (3: 1), narrowly lanceolate (6: I) to narrowly ovate (2: 1) or narrowly 
oblong (3: 1), 4 to 21 cm. long including the stipes, 1 to 4 cm. (rarely up to 6-8 cm.) 
broad, internal cells with thick, pitted walls, the apex erect or deflexed, often caudate. 
Primary pinnae petiolulate, 7 to 17 pairs, rather widely spaced towards the base of 
the frond, lanceolate, rhomboidal to flabellate, 0-5 to 4 7 cm. long, 0-5 to T2 cm. 
broad, the apex acute or caudate. Ultimate segments sessile, linear (12: 1) to 
narrowly oblong (3: 1) but mostly cultrate (6: 1), 1 to 5 mm. long, 0-2 to 0-5 mm. 
broad, simple or bifid, 1-nerved, glabrous or with a few, scattered hairs, the apex 
obtuse, truncate or emarginate. Veins dark, prominent, not reaching the margin, 
with a few, 2- to 4-celled, sometimes glandular-tipped hairs. Sori indusiate, 12 to 
130 per frond, 1 to 11 per primary pinna, sunken in an inner basal lobe of a secondary 
pinna. Involucre 0-5 to 1 mm. long, 0-5 to 0-8 mm. broad at the apex, usually very 
narrowly winged on both sides, cupuliform to almost triangular, the base cuneate, 
the border or mouth spreading or almost imperceptibly expanded, not reflexed, 
often 2-lipped, the margin entire or slightly erose or sometimes fluted. Receptacle 
extruded, projecting 1 to T5 mm. beyond the border. Sporangia subsessile, 
compressed, the oblique annulus composed of 16 to 21 indurated cells and 6 to 8 
thin-walled cells, dehiscing by a lateral slit. Spores globose-tetrahedral to 
globose, trilete, the laesurae long, low ridges, the exine thin, suberanulate, equat. 
diam. 24-38 to 31-88^ x polar diam. 24-38 to 31-88g. (No. 201, PI. VI, f. 3 and 4.) 


44 


201 . Hymenophyllaceae. (Tin dole) 


Distribution: Tahiti, Samoa, Fiji, New Caledonia and Australia (north-eastern 
Queensland to Victoria). 

Holotype: Tahiti, Society Islands, Wilkes (US 51176), examined. 

Queensland: Herberton, Waller No. All, 1909 (NSW P2130); Fraser Island, near the 
Forestry Station, epiphytic on Dicksonia youngiae at the edge of a creek, rhizome creeping, 
fronds horizontal to pendulous, Blake No. 14372, 8.1941 (BRI); Fraser Islands, Wide Bay, 
White 10.1921 (BRI); Eumundi, Simmonds (BRI); Buderim Mountain, White 4.1916 (BRI; 
NSW P2128); Wide Bay District, Montville, in rainforest on tree-fern, ca. 600 ft. alt., rhizome 
creeping, fronds light green, Blake No. 13836, 8.1938 (BRI); Tambourine Mountain, Simmonds 
10.1909 (NSW P2127); Tallebudgera Creek, Schneider (BRI); Nerang River, Schneider (BRI); 
Roberts Plateau, Lamington National Park, exceedingly common on tree trunks, fronds pale 
green. Smith & Goy No. 70, 1.1938 (BRI; K); Binna Burra, 2,600 ft. ait., Dickson 9.1947 
(NSW P5329); Lamington National Park, in rainforest, 3,500 ft. alt., Blake No. 12987, 5.1937 
(BRI); Springbrook, on trunk of tree-fern. White 12.1915 (BRI); Macpherson Range, Tryon 
3.1891 (BRI). 

New South Wales: Whian Whian State Forest, WNW of Byron Bay, 2,050 ft. alt., 
pendulous on the trunks of tree-ferns, frequent in rocky gully, rainforest, basalt, Constable 
1.1953 (NSW P6483); Tuckombil, Alstonville, Watts 11.1902 (NSW P1147, P1142); Uralba, 
on trunk of tree-fern, Bauerlcn No. 272, 5.1891 (NSW PI 863); East Bailina, on tree-ferns, 
Watts No. 55, 4.1899 (NSW P1143); Three Mile Scrub, Richmond River, Watts 11.1902 (NSW 
PI 144); Richmond River, Watts 1901 (NSW PI 146); Dorrigo, Heron 4.1910 (NSW P2129); 
Bcllenger River, Moore (NSW P2133); Port Macquarie, Moore, received 5.1867 (K); William’s 
River, Fraser & Vickery 1.1934 (NSW P2126); Ourimbah, near Wyong, Helms 10.1904 (NSW 
P583); Gosford, in rainforest, Deane 10.1888 (NSW PI 122); Valley of the Waters, Watts 9.1912 
(NSW PI 145); Illawarra, Moore (NSW P2131); £ mile NE of Trig., Mt. Dromedary, Central 
Tilba, 2,100 ft. alt., frequent on tree-fern trunks in rainforest, Constable 9.1953 (NSW P6641). 

Victoria: Gembrook, Lucas 11.1876 (NSW P8281), 

Absolute Synonym: Trichomanes caudatum Brack., Ferns U.S. Expl. Exped. XVI (1854) 
256, t. 36, f. 5; Benth., FI. Austr. VII (1878) 703; Bail., Lith. Ferns Qucensl. (1892) 28; Moore 
& Betche, Handb. FI. N.S.W. (1893) 503; Maiden & Betche, Census N.S.W. PI. (1916) 1; 
Copel. in B.P. Bish. Mus. Bull. No. 93 (1932) 24; Copel. in Phil. Journ. Sci. LI (1933) 262, pi. 
57, f. 3-5, pi. 58, f. 1; C. Christensen in B.P. Bish. Mus. Bull. No. 177 (1943) 18. 

The Australian material is undoubtedly a geographic segregate of M. caudata but whether 
it should be treated as a subspecies or as a new species would depend on chromosome counts 
of M. caudata from Australia as well as from Tahiti, Samoa, Fiji etc. The New Caledonian 
materia] which was described by van den Bosch as Trichomanes tnilnci in Ann. Sci. Nat. IV, 

15 (1861) 89, is more distinct and should probably be treated as a separate species. The holotype 
which was collected by Vieillard on the trunks of tree-ferns at Balade, was examined at the 
Museum d’Histoire Naturelle, Paris, by the author. 

An anatomical examination of transverse sections of the stipes of M. caudata collected 
in Fiji (Parks No. 20770), New Caledonia (NSW P7297) and Australia (NSW P8282) revealed 
only slight differences in structure. Under a single row of thick-walled epidermal cells there 
are 3 to 5 rows of sclerenchyma enclosing about 5 rows of collenchyma which surround the 
central vascular bundle. 

The Australian material is characterized by smaller, often truncate, narrower involucres 
(i.e. 0-5 to 1 mm. long and 0-5 to 0-8 mm. broad at the apex) than in the specimens from any 
of the Pacific Islands. The Fijian material (which approximates most closely to the Australian) 
has involucres which range from 1 to 1 5 mm. long and 0-5 to 1 •2 mm. broad at the apex, e.g. 
A. C. Smith No. 5106 (US) collected at Mba, Viti Levu, Fiji, has particularly small involucres. 
The involucres of the Samoan specimens are 1 -2 to 2 mm. long whereas those from Tahiti range 
from 1 to 1-5 mm. long and 0 5 to 0-9 mm. broad. Setchell & Parks No. 538 (US) collected 
“ on the face of the Diadem, Tahiti ”, in July. 1922, is characterized by very small sori which 
are 1 to 1 -2 mm. long and 0-5 to 0-8 mm. broad at the apex. In general appearance it is similar 
to Eastern Australian material. The Holotype of M. caudata (i.e. US 51176), which was 
collected at Tahiti, has much larger involucres, being 1-3 to 1-5 mm. long and 0-8 to 0-9 mm. 
broad at the apex. In the New Caledonian material of M. caudata, which was described as 
Trichomanes milnei by van den Bosch in Ann. Sci. Nat., IV, 15 (1861) 89, the mouth of each 
involucre is more dilated than in the Australian specimens, also the involucre is much longer, 
i.e. 1 *7 to 2 mm. in length. 


201. Hymenophyllaceae. ( Tindale ) 


45 


The fronds of the Australian specimens tend to be smaller than in the Pacific Island 
material, i.e. 4 to 21 cm. long including the stipes, whereas the Fijian and Tahitian material is 
up to 35 cm. long. 


10. CEPHALOMANES Presl 

Cephalomanes Presl, Hymen. (1843) 17, pi. 5; Copel. in Phil. Journ. Sci. 
LXV1I (1938) 66; Copel., Gen. Fil. (1947) 40. Trichomanes L. subgen. Cephalomanes 
(Pr.) C. Chr., Ind. Fil., Suppl. 3 (1934) 187; Holttum, Ferns of Malaya (1954) 102. 

In Phil. Journ. Sci. LI (1933) 245, Copeland treated Cephalomanes as a group of 
Trichomanes L. 

Terrestrial ferns. Rhizome erect, tufted, with thick roots which become 
stilt-like in age. Stipes fascicled, bearing stiff, dark red hairs when young. Fronds 
caespitose. Lamina 1-pinnate or rarely 1-pinnatifid, dark green, harsh, setose, 
polystichous, lanceolate, more or less elliptical, cultrate or oblong, the cell-walls 
thin and wavy, the cells large and with dark contents. Pinnae overlapping, unequally- 
sided, the upper side more or less lacerate and the lower with long, prong-like, 
upcurved teeth. Veins coarse, rib-like, forked, often clothed with bulbous, unicellular 
hairs. Sori numerous, mostly borne on the upper pinnae, terminal on the veinlets 
on the acroscopic side, rarely reaching the basiscopic side. Involucre cylindrical or 
rarely obconic, thick. Receptacle long-extruded, thick, sometimes with an enlarged 
apex. Sporangia subsessile, compressed-turbinate, with a very oblique annulus. 
Spores tetrahedral or globose-tetrahedral to almost oblong, trilete, green. 

Type of Genus: Cephalomanes atrovirens Presl, native to the Philippines, 
New Guinea and Australia. 

This is a very distinctive genus of about 10 species ranging from Madagascar 
to India and Polynesia. According to Copeland in Genera Filicum (1947) 40, it 
is related to Vandenboschia especially to V. auriculata. There is only one Australian 
species of Cephalomanes, viz. C. atrovirens which ranges from the tropical rainforests 
of north-eastern Queensland to the subtropical North Coast of New South Wales. 

A chromosome count of n = ca. 32 was made by Manton on Malayan 
material of Cephalomanes javanicum (Bl.) v.d. Bosch (as Trichomanes javanicum Bl.) 
in the Appendix to Holttum, Ferns of Malaya (1954) 623. 


Cephalomanes atrovirens Presl, Hymen. (1843) 18 t. 5; Copel. in Phil. 
Journ. Sci. LXVII (1938) 67. 

Habitat terrestrial in tropical and subtropical rainforests, often on creek 
banks or sometimes in wet soil not far from a stream. Rhizome erect, tufted, the 
roots stick-like, woody and 1 to T5 mm. in diam. Stipes not winged, rather short, 
about one fifth to one eighth the length of the frond, 1 to 4-5 cm. long, fasciculate, 
flattened towards the apex and with 1 or 2 grooves on the upper surface, terete 
towards the base, brown, woody, dull, scabrous, sparsely clothed with scattered, 
spreading, glossy, elongated, dark red, articulated hairs. Main rhachis very narrowly 
winged except at the base, thick, woody, flattened broadly, 1-grooved and glabrous 
above, densely clothed on the lower surface with crisped, acute, dark red hairs of 
the same type as on the stipes. Lamina 1-pinnate, herbaceous to subcoriaceous, 
harsh, dark green, compact, very narrowly elliptical (6: 1) to elliptical (2: 1), cultrate 
(6: 1) to oblong (2: 1), 10 to 23 cm. long, 1 to 3-7 cm. broad, often very much 


46 


201. Hymenophyllaceae. (Tindale) 


decurved, not dimorphic, the apex usually acute, the base mostly tapering, i.e. the 
lower pairs of pinnae shorter; internal cells of the lamina comparatively thin and 
wavy, the cells large with dark contents. Primary pinnae with short, winged, 
persistent petiolules, 20 to 45 pairs, imbricate, narrowly oblong (3: 1) to oblong 
(2: 1), 1 to 2 cm. long, 2-5 to 6 mm. broad, the apex broadly rounded or obtuse, 
the base very unequally-sided, the upper side truncate, the lower margin much 
narrowed and cut away, the acroscopic margin laciniate-dentate, the basiscopic 
margin with long, upcurved, prong-like laciniae on the adaxial surface. Veins 
forked, dark green, prominent, raised above the lower laminal surface, rib-like, 
much branched, parallel from the costa and reaching the margin, produced into 
short, setaceous teeth on the acroscopic margin and into long laciniae on the basiscopic 
margin, the surface of the veins clothed with fawn, unicellular, appressed, oblong- 
bulbous glandular hairs. Sori indusiate, marginal, 1 to 6 on the upper margin 
of each pinna, confined to the proximal half or each primary pinna, usually 60 to 
200 on each frond, occurring throughout the lamina, free, almost wholly exserted, 
often subtended by a marginal tooth, narrowly winged, terminal on a minor veinlet. 
Involucre 1 to 2 mm. long, 0-5 to 0-8 mm. broad at the middle, firm, tubular, narrow 
and elongated, dark green or brown, clothed with small, fawn, unicellular, oblong- 
bulbous, appressed, glandular hairs, the small rim glabrous, slightly dilated, the 
base narrowly cuneate. Receptacle long-extruded, coarse, projecting 1 to 6 mm. 
beyond the mouth of the involucre, the tip mostly thickened and knob-like. 
Sporangia subsessile, compressed-turbinate, the annulus very oblique, consisting of 
15 to 18 indurated cells and about 3 thin-walled cells. Spores subtriangular to 
almost oblong, trilete (the laesurae broad), granulate, equat. diam. 31-88 to 39-38g 
X polar diam. 24-48 to 30 |a. (No. 201, PI. VI, f. 5 and 6.) 

Distribution: The Philippines, New Guinea and Australia (north-eastern Queensland 
to the North Coast of New South Wales). 

Holotype: Insulas Philippinas, Cuming No. 169 (K), examined. 

Queensland: Cooktown Tableland, on wet rocks in rainforest near creek, Messmer 
7.1952 (NSW P6379); Daintree River, Fitzalan. ex Herb. Muell. 7.1877 (K); ditto, Pentzcke 
1881 (NSW P4988); ditto, on wet soil in rainforests. Brass No. 2209, 2.1932 (BRI); ditto, in 
damp heavy soil in rainforest gullies, Messmer 7.1954 (NSW P6876); ditto, growing on rocks 
along creek banks close to the water, Messmer 8.1952 (NSW P2343); Black Mountain Rd.- 
Kuranda, on creek banks, almost in the water, in rainforest, Messmer 8.1952 (NSW P2333); 
Kuranda, Waller 12.1908 (NSW P2138) and 1908 (NSW PI 159); in a gully at the foot of Smithfield 
Range, Kamerunga. near Cairns, Cowley 7.1890 (BRI); West Cairns Hills, 1,000 ft. alt., in wet 
rock crevices in rainforest gorge, Messmer 6.1954 (NSW P6877); Cairns, Wild 1891 (BRI); 
between Cairns and Herberton, Wild 1891 (BRI); Johnstone River, Kefford (BRI). 

Absolute Synonym: Trichomanes atrovirens (Pr.) Kunze in Bot. Zeit. V (1847) 371; 
Copel. in Phil. Journ. Sci. LI (1933) 251, t. 52 f. 3, t. 55 f. 2; C. Chr., Ind. Fil., Suppl. 3 (1934) 


Equivalent Synonym: Trichomanes rhomboideum J. Sm. in Hook. Journ. Bot. Ill 
(1841) 417, nomen nudum, Holotype by Monotypy: Luzon (the Philippines), H. Cuming 
No. 169 (K), examined. J. Smith stated that this specimen was located at the Kew Herbarium. 

This species was misidentified as Trichomanes javanicum auett. non Bl. by the following 
authors:—Bcnth., FI. Austr. VII (1878) 702); F. M. Bail., Lith. Qucensl. Ferns (1892) t. 24; 
Domin in Bibl. Bot., LXXXV0913) 13; Maiden & Betche, Census N.S.W. PI. (1916) 1; Melvaine 
in Proc. Linn. Soc. N.S.W. LXI (1936) 114. 

The record of Cephalomanes atrovirens for the North Coast of New South Wales is 
somewhat doubtful. However, several ferns and fern-allies have been recorded for this State 
after intervals of 40 to 60 years, so that this species is included for completeness. 

The Australian specimens of C. atrovirens agree well with material of this species collected 
in the Philippines. It is closely allied to C. javanicum v.d. Bosch which ranges from Burma 


201 . Hymenophyllaceae. ( Tindale ) 


47 


to Malaysia to New Guinea but is separated from the latter species by the slightly dilated (instead 
of truncate) involucres and the shorter stipes. In C. atrovirens the stipes are 1 to 4-5 cm. in 
length whereas they are 8 to 12 cm. long in C. javanicum. As indicated by Copeland in Phil. 
Journ. Sci. LI (1933) 251-2, C. atrovirens is distinguished by its “ narrow, very compact fronds 
with very numerous, small, imbricate pinnae, dark colour and thin texture ”. 

C. atrovirens belongs to the group of species in which the sori are borne on normal pinnae 
not in a terminal spike or panicle. 


11. SELENODESMIUM (Prantl) Copel. 

Selenodesmium Prantl, Hymen. (1875) 53, as a section of Trichomanes; 
Copeland in Phil. Journ. Sci. LI (1933) 232, as the group of Trichomanes rigidum-, 
Holttum, Ferns of Malaya (1954) 108, as a subgenus but the combination was not 
made; Copeland in Phil. Journ. Sci. LXVII (1938) 80 and Gen. Fil. (1947) 42, as 
a genus. 

Terrestrial or rarely rupestral ferns. Rhizome erect, tufted or shortly creeping, 
clothed with dark, bristle-like hairs and bearing stout roots which occasionally 
become stilt-like in age. Stipes usually more or less fascicled, not winged, elongated, 
clothed with dark, deciduous, bristle-like hairs which are under 2 mm. long. Lamina 
usually ovate to deltoid, finely decompound, harsh or firm in texture without false 
veins or thickened margins, the cell-walls coarsely pitted, thickened and straight or 
wavy. Pinnules deeply dissected but with a broad, uncut, medial portion. Ultimate 
segments narrow. Veins closely parallel, then diverging into the segments. Involucre 
cylindrical, the mouth entire or slightly 2-lipped. Receptacle usually long-exserted. 
Sporangia subsessile, small and compressed. Spores tetrahedral-globose to globose, 
trilete, scabrate. 

Type of Genus: Selenodesmium rigidum (Sw.) Copel. (Basionym: Trichomanes 
rigidum Sw.), native to Jamaica. 

This is a small, well-defined genus of about 10 species ranging from Formosa 
and India to Australia and New Zealand through Polynesia to Jamaica. It is 
mainly found in the tropical and subtropical parts of the world. In Australia 
only one species has been recorded, viz. Selenodesmium elongatum (A. Cunn.) 
Copel. which occurs in Queensland and on the North Coast of New South Wales. 

The chief diagnostic characteristics of the genus are the deltoid fronds and 
the thick, coarsely pitted cell-walls of the lamina. This is an essentially terrestrial 
or rarely rupestral genus of ferns which prefer dark, shady forest gullies. Epiphytic 
hepatics commonly grow on the laminas of Selenodesmium spp. 

A chromosome count of n = 33 was recorded for Selenodesmium obscurum 
(Bl.) Copel. (as Trichomanes obscurum Bl.) by Manton & Sledge in Phil. Trans. 
Roy. Soc. London, Ser. B, Biol. Sci. No. 654, Vol. CCXXXVIII (1954) 136, from 
material collected in Ceylon. They also recorded a count of n = ca. 33 for Malayan 
specimens of the same species, l.c. 136. 


Selenodesmium elongatum (A. Cunn.) Copel. in Phil. Journ. Sci. LXVII (1938) 
92; Dobbie & Crookes, New Zealand Ferns (1952?) 118, 119 (photo.). 

Habitat terrestrial or sometimes rupestral in tropical or subtropical rainforests, 
in damp gullies along creek banks or sometimes under rock ledges. Rhizome short, 
thick, erect, oblique or very shortly creeping, 0-7 to 2 cm. high, densely clothed with 


48 


201 . Hymenophyllaceae. ( Tindale) 


simple, septate, almost black, acute hairs which are 1 to 2 mm. long, the roots woody, 
prop-like and rigid. Stipes not winged, almost terete, with 2 insignificant longitudinal 
ridges, fascicled, 3-5 to 20 cm. long, 0-5 to 1 mm. broad, terete, rigid, brown or 
dark purplish-brown, darker on the adaxial surface, woody, smooth or slightly 
scabrous, glabrous or bearing a few, almost black, deciduous, fibrillose hairs near 
the base. Main rhachis very narrowly winged towards the apex by the decurrent 
bases of the primary pinnae. Fronds 8 to 36 cm. long including the stipes. Lamina 
3-pinnate or 3-pinnate-pinnatifid, thinly coriaceous, dark olive-green, rigid, lanceolate- 
deltoid (3: 1) to ovate-deltoid (3: 2), 5 to 14-5 cm. long, 2 to 9 cm. broad, the cell-walls 
very thick and pitted, not wavy, the apex acute to acuminate, either erect or deflexed. 
Primary pinnae petiolulate, opposite at the base but alternate towards the middle 
and apex, narrowly ovate-deltoid (2: 1) to broadly ovate-deltoid (6: 5), 1 -5 to 4-5 cm. 
long, 1 to 2-5 cm. broad, the apex obtuse to acute, seldom acuminate. Secondary 
pinnae with unwinged petiolules except near the apex of each primary pinna, much 
dissected at the base, the uncut lamina oblanceolate; apex obtuse, subacute or 
broadly rounded, finely denticulate. Ultimate segments flat, usually oblanceolate 
(3: 1) to narrowly obovate (2:1) or cultrate (6: 1), denticulate towards the apex. 
Veins prominent, almost parallel then diverging into the ultimate segments and 
much forked dichotomously, reaching the margin, densely clothed on the lower 
surface with minute, scurfy, yellowish-brown, 1- to 3-celled hairs. Sori 20 to 200 
on each frond, 1 to 18 on each primary pinna (occurring on both margins), borne 
on short lobes in the axils of the tertiary segments. Involucre 1 -5 to 2 mm. long, 
the middle of the tube 0-5 to 0-6 mm. broad, cylindrical, sometimes reflexed, dark 
olive-green or brown, free but narrowly winged on both sides throughout its length, 
clothed with scurfy hairs especially near the base; mouth 0-5 to 0-8 mm. broad, 
truncate or slightly dilated and reflexed, entire or incipiently 2-lipped; base narrowly 
cuneate, greatly attenuated. Receptacle very long-exserted, curved, projecting up 
to 1-5 cm. beyond the mouth of the involucre, the apex unthickened or slightly 
dilated. Sporangia subsessile, compressed, the annulus obliquely transverse, 
composed of 15 or 16 indurated cells and about 4 thin-walled cells. Spores globose- 
tetrahedral to globose, trilete, white (later green), the laesurae simple, the exine 
very thin and scabrate, equat. diam. 28-25 to 31 -88;x X polar diam. 22-5 to 28-13g. 
(No. 201, PI. VI, f. 1 and 2.) 


Distribution: New Zealand (North and South Islands) and Australia (North-eastern 
Queensland to the North Coast of New South Wales). 

Holotype: In damp woods of Wangaroa and Hokianga No. 56, R. Cunningham, 1834 
(K), examined. This specimen was collected in New Zealand. 


Queensland: Cooktown Tableland, growing on rocks in the bed of a river, in rainforest, 
Messmer 7.1952 (NSW P4871); Cooktown, Harris (BRI 30339); Thornton Peak, alt. 4,000 ft., 
Brass & White No. 228, 9.1937 (BRI 30372); Daintree River, opposite Mission, near creek, 
in leaf mould in damp gully, Messmer 7.1954 (NSW P6824); dillo, in rainforest, Messmer 
8 1952 (NSW P22I8); Mt. Demi (4 miles SW of Mossman, 3.000 ft. alt., on wet mountain slopes, 
Brass No. 2082, 2.1932 (BRI 30366-7); Black Mountain Road-Kuranda, in rainforest, on creek 
banks, almost in water, Messmer 8.1952 (NSW P2332); Street’s Gully, Kuranda, Watts 7.1913 
(NSW P3759); West Cairns Range, 1,000 ft. alt., under rock ledges in rainforest gorges, Messmer 
7.1954 (NSW P6875); Cairns district. Watts 7-8.1913 (NSW P2092); Cairns, Froggatt 1886-7 
(NSW PI 148); Atherton, McKenzie 9.1910 (NSW P2096); Atherton Scrub Mitchell 8.1911 
(NSW P209I); Bellenden Ker, Hamilton Kenny 1.1912 (BRI 30342); Malanda,^tufted, oblique 
to erect, dark green, on banks in rainforest, 2,400 ft. alt.. Blake No. 15171, 8.1943 (BRI 30369), 
Herberton. Waller 1908 (NSW P2095); Johnstone River, Kcfford (BRI 30336); ditto Bancroft 
(BRI 30340); creek on the way to Tullv Falls, Watts 7-8.1913 (NSW P3758); between Cleveland 
Bay and Rockingham Bay, Hill Nos. 57 and 58, 1.1866 (K); Seaview Range Temson-Woods 
1875 (NSW P2094); Paluma Range, in rainforest. Kennedy 8.1948 (NSW P4988); Dalrymple 
Heights, on stream bank in forest, Clemens 8-11.1947 (BRI 20187); Eumundi, Simmonds 11.1892 
(K; BRI 30363-4); Yandina, Bailey 3.1891 (NSW P582); ditto, Field Naturalists 5.1891 (BRI 
30355); Maroochie, Bailey 7.1879 (BRI 30337); Buderim Mountain, White 4.1916 (BRI 3;34)30 


201 . Hymenophyllaceae. ( Tindale ) 


49 


Eudlo Creek, Simmonds (BRI 30356); Pearson’s Falls, Helidon, Bailey 10.1883 (BRI 30338); 
Nerang Creek, Schneider (BRI 30341). 

New South Wales: Macpherson Range, Campbell 8.1901 (NSW P2093); Main Arm 
of the Brunswick River, near Mullumbimby, fairly common on shady creek banks in wet alluvial 
soil, in rainforest, Gov No. 135, 8.1936 (BRI 30365); Tintenbar. Bauerlen 12.1896 (NSW PI856)- 
Richmond River, Watts 1901 (NSW P3757); Clarence River (NSW P2097). 

Absolute Synonyms: Trichomanes elongatum A. Cunn., Comp. Bot. Mag., II (1837) 
368; Hook., Ic. PI. (1845) t. 701; Copel. in Phil. Journ. Sci. LI (1933) 239, pi. 47, f. 1 and 2. 
Trichomanes rigidum Sw. var. elongatum (A. Cunn.) Hook, et Bak. in Hook. & Bak., Syn. Fil. 
(1873) 86; Thomson, The Ferns & Fern Allies of New Zealand (1882) 48, 

Equivalent Synonym: Trichomanes polyodon Col. in Trans. N.Z. Inst. XXVIII (1896) 
618, Isotype: New Zealand, presented by W. Colenso in 1897 (K). 

This species was misidentified as Trichomanes rigidum Sw. in the following publications:— 
Benth., FI. Austr. VII (1878) 702; Moore & Betche, Handb. FI. N.S.W. (1893) 503; Maiden 
& Betche, Census N.S.W. PI. (1916) 1; Melvaine in Proc. Linn. Soc. N.S.W. LXI (1936) 114. 
However, Selenodesmium rigidum (Sw.) Copel. (syn. T. rigidum Sw.) is a native of Tropical 
America. 

Specimens such as NSW P2095 collected by Waller at Herberton, North Queensland, 
are a good match for material of Selenodesmium elongatum obtained in New Zealand, e.g. BRI 
30349 from Nelson and AK 31491 from Walkers Bush, Auckland. Nevertheless, the laminas 
of these specimens are less finely dissected than in most Australian plants of this species. S'. 
elongatum was originally described from New Zealand where it is found in both the North and 
South Islands on wet, shady banks from sea level to 2,500 ft. altitude according to Harris, Spores 
of New Zealand Pteridophyta (1955) 84. 

There is a good deal of variation in the Australian material of S. elongatum especially 
in the degree of dissection of the secondary pinnae. There is every gradation between specimens 
such as NSW P2095 from Herberton where the secondary pinnae are shallowly cut into short 
broad lobes, to NSW P6824 from the Daintree River. In the latter frond these secondary 
pinnae are very finely dissected into narrow segments. 

S. elongatum differs from S. obscurant (Bl.) Copel. which is common in Ceylon, Malaya, 
Southern India, Formosa, Indonesia and New Guinea, in the usually uncurled tips of the laminal 
segments in the former species. This point is difficult to assess in dried specimens because of 
the number of badly pressed fronds but these tips are uncurled in living plants which I have 
examined. The fronds are much more finely dissected in S. obscurant, also the uncut lamina 
of the pinnules is mostly linear instead of oblanceolate or obovate as in S. elongatum. Similarly 
in another closely allied species viz. S. dentatum (v.d. Bosch) Copel., of Polynesia, it is linear 
to elliptical (see Copeland in Phil. Journ. Sci. LI (1933) 233), but the tips of the laminal segments 
are uncurled as in S. elongatum. 

Hepatics are commonly found growing on the laminas of S. elongatum, e.g. Lepidozia 
gottscheana Ldgb. is epiphytic on NSW P4871 collected by Messmer on the Cooktown Tableland, 
North Queensland. Miss Helen Hewson, University of Sydney, kindly made the identification 
of this bryophyte for me. 

In Eastern Australia this terrestrial (or less commonly rupestral) fern is mostly found 
on shady, wet creek banks in good alluvial soil. S. elongatum occurs from the tropical rainforests 
of north-eastern Queensland to the subtropical rainforests of the Far North Coast of New South 
Wales but it has not been recorded from Nothofagus forests. 





No. 201, Plate 1 


M. D. Tfndale del. 



1. Sphaerocionium lyallii , habit study, X 2$. 2. 5. lyallii, sorus at apex of an ultimate 

segment, x 18. 3. Apteropteris malingii, stellate laminal hair, X ca. 100. 4. A. malingii, sorus on 

portion of primary pinna, x 18 . 


G99777 5 
















No. 201, Plate II 


M. D. Tindale del. 



Hymenophyllum spp. 


H. marginatum, habit study, X 4£. 
3. H. rarum, portion of fertile frond, x 5. 


2. H. marginatum, sorus at apex of lamina, x 15. 
4. H. cupressiforme, portion of fertile frond, x 5. 











No. 201, Plate III 


M. D. Tindale del. 



Hymenophyllum flabellatum 

I. Large fertile frond, x ^ natural size. 2. Two sori on ultimate segments, X 4. 3. Sorus 

with receptacle bearing sporangia, x 33. 4. Small sterile frond on rhizome, x 5. Spore, 

X 320. 6. Hair of rhizome, ca. X 40. 




















No. 210, Plate IV 


M. D. Tindale del. 



Polyphlebium venosum 


I. Lower portion of frond with a fully developed primary pinna and several reduced 
pinnae, X 5. 2. Sporangium dehiscing with 2 spores, x 140. 3. Sorus with cylindrical involucre, 

x 25. 4. Spore, X 640. 5. Cells of lamina, x 250. 
























No. 201, Plate V 


M. D. Tindale del. 



2 


3 


Dwarf Species 


1. Microtnchomanes vitieiise , habit study x 44 
study, x 4J. 3. MicrogOnium tahitense, habit study, x 


2. Microgonium bimarginatum , habit 
6 $. 


G 99777—6 



















No. 201, Plate VI 


M. D. Tindale del. 




6 


The Tricliomanes Group 

1. Selenodesmium elongawm, portion of a primary pinna with sori, x 3T 2. S. elongation, 
cells of lamina with pitted walls, x ca. 266. 3. Macroglena caudata, cells of lamina with pitted 

walls, x ca. 266. 4. M. caudata , primary pinna with sori, x 2}. 5. Cephalometries atrovirens, 

primary pinna with sori, x 3T 6. C. atrovirens, cells of lamina with pitted walls, x ca. 266. 



























No. 201, Plate VII 


Photograph by J. Peacock 



del. M. Tindale 



Hymenophyllum peltatum 

Above: Mitotic chromosomes, late prophase, X 900. Below: Diagram of the same cell, 
X 1,800. 


93 Crassulaceae 

94 Escalloniaceae 

95 Pittosporaceae 

96 Cunoniaceae 

97 Davidsoniaceae 

98 Rosaceae 

99 Mimosaceae 

100 Caesalpiniaceae 

101 Papilionaceae, Part 

102 Geraniaceae 

103 Oxalidaceae 

104 Linaceae 

105 Erythroxylaceae 

106 Zygophyllaceae 

107 Rutaceae 

108 Simaroubaceae 

109 Burseraceae 

110 Meliaceae 

111 Tremandraceae 

112 Polygalaceae 

113 Euphorbiaceae 

114 Cailitricbaceae 

115 Anacardiaceae 

116 Aquifoliaceae 

117 Celastraceae 

118 Siphonodontaceae 

119 Hippocrateaceae 

120 Stackhousiaceae 

121 Icacinaceae 

122 Sapindaceae 

123 Akaniaceae 

124 Rhamnaceae 

125 Vitaceae 

126 Elaeocarpaceae 

127 Tiliaceae 

128 Malvaceae 

129 Sterculiaceae 

130 Dilleniaceae 

131 Eucryphiaceae 

132 Hypericaceae 

133 Elatinaceae 

134 Frankeniaceae 

135 Violaceae 

136 Flacourtiaceae 

137 Passifloraceae 

138 Cactaceae 

139 Thymelaeaceae 

140 Lythraceae 

141 Rhizophoraceae 

142 Myrtaceae 


143 Melastomataceae 

144 Onagraceae 

145 Haloragiaceae 

146 Araliaceae 

147 Umbclliferae 

148 Alangiaceae 

149 Ericaceae 

150 Epacridaceae 

151 Myrsinaceae 

152 Primulaceae 

153 Plumbaginaceae 

154 Sapotaceae 

155 Ebenaceae 

156 Symplocaceae 

157 Oleaceae 

158 Loganiaceae 

159 Gentianaceae 

160 Menyanthaceae 

161 Apocynaceae 

162 Asclcpiadaceae 

163 Convolvulaceae 

164 Polemoniaceae 

165 Hydrophyllaceae 

166 Boraginaceae 

167 Verbenaceae 

168 Avicenniaceae 

169 Labiatae 

170 Solanaceae 

171 Scrophulariaceae 

172 Selaginaceae 

173 Bignoniaceae 

174 Pedaliaceae 

175 Martyniaceae 

176 Orobanchaceae 

177 Gesneriaceae 

178 Lentibulariaceae 

179 Acanthaceae 

180 Myoporaceae 

181 Plantaginaceae 

182 Rubiaceae 

183 Caprifoliaceae 

184 Dipsacaceae 

185 Cucurbitaceae 

186 Campanulaceae 

187 Lobeliaceae 

188 Goodeniaceae 

189 Brunoniaceae 

190 Stylidiaceae 

191 Compositae 


PTEIUDOPHYTA 


192 Lycopodiaceae 

193 Selaginellaceae 

194 Isoetaceae 

195 Psilotaceae 

196 Ophioglossaceae 

197 Maratliaceae 

198 Osmundaceae 

199 Schizaeaceae 

200 Gleicheniaceae 

201 Hymenophyllaceae 

202 Cyatheaceae 

203 Dicksoniaceae 

204 Lindsaeaceae 


205 Pleridaceae 

206 Adiantaceae 

207 Vittariaceae 

208 Davalliaceae 

209 ydrammitidaceae 

210 Polypodiaceae 

211 Aspidiaceae 

212 Thelypteridaceae 

213 Aspleniaceae 

214 Blechnaceae 

215 Marsileaceae 

216 Azollaceae 

217 Salviniaceae