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Volume 74, 1944-45
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New Zealand Hepaticae (Liverworts).—IV.
A Review of the New Zealand Species of the Genus Radula

[Read before Wellington Branch, August 23, 1944; received by the Editor, August 23, 1944; issued separately, December, 1944.]

The much discussed question as to why Dumortier's name of Radula should have been retained in preference to Gray's earlier name of Martinellius, is fully dealt with in Dr. H. Castle's “Revision of the Genus Radula, 1, 1936.” The primary reason for the ignoring of the name Martinellius was the masculine termination, and when Carruthers drew attention to it in 1865, the name Radula was too firmly established to be altered. But there is much more to it than that. Other plants now recognised as belonging to other genera were included in Martinellius, thus involving more than one application of what is known as the “method of residues,” with the final result that the name Radula “descends to the genus which has borne that name with nearly every hepaticologist since 1833.”

Genus Radula Dumortier.

Radula Dum., Comm. Bot., 112, 1822; Rec. d'Obs., 1835.

Martinellius S. F. Gray, Nat. Arr. Brit. Pl., 1, 690, 1821.

Stephanina O. Kuntz, Rev. Gen. Pl., 839, 1891. Schiffn. in Engl. & Prantl, Nat. Planfz., 1895.

Plants usually dioicous, may be autoicous or paroicous, medium, rarely small, in various shades of green, from yellowish to dark, often imparting a yellow-green stain to paper when wet, usually creeping and forming depressed mats on bark, earth, rock, mosses or filmy ferns. Stems usually pinnately to bi-pinnately branched, branches infrafoliar in origin. Leaves incubous, alternate, entire or toothed, complicate-bilobed; ventral lobe smaller, appressed to the underside of the dorsal lobe, or with the carinal portion inflated, rhizoids often arising from a mamilliform protuberance. Stipules everywhere absent. Involucral leaves usually larger than the foliage. Perianths usually acrogynous, frequently with 1–2 floral innovations, dorsiventrally compressed, plicate or smooth, oblong or with an elongated neck; lips truncate, crenate, undulate, or with a median cleft, margins sometimes winged. Capsule generally oval-cylindrical, valves with 2 layers of cells, dehiscing to the base. Androecia terminal or intercalary, bracts in pairs from 3 to about 30, antheridia usually single. Vegetative reproductive bodies present or absent.

A fairly large genus, with numerous tropical species.

Castle (1937) has divided the sub-genus Acroradula Spruce into 13 sections, using where fitting, the limitations and names of 4 of Stephani's 7 sub-genera, and regrouping and renaming more satisfactorily, the remaining 3.

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The distinguishing features of the sub-genera and sections are as follows, the following species having been placed in their respective sections on the authority of the writer:—R. papulosa, sainsburiana, buccinifera, plicata, levieri, silvosa, allisonii.

Subgenus Cladoradula Spruce.

Perianth terminal on short lateral branches only.

R. physoloba (sterile in New Zealand).

Subgenus Acroradula Spruce.

Perianth terminal on the main stem or on leading branches; sub-floral innovations may cause it to be falsely lateral.

Section Amentulosae St.

Microphyllous branches present.

R. uvifera.

Section Ampliatae St.

Upper half of the lobule base free, rounded, and extended well across or beyond the stem.

R. grandis.

Section Complanatae Castle.

“… Basal portion of the lobule free for less than one-half its length and extended usually less than half the distance across the axis.”

R. papulosa.

Section Adnatae Castle.

“… Base of the lobule fused to the axis for practically its entire length.”

R. sainsburiana, buccinifera, plicata, levieri, silvosa.

Section Longilobae St.

Lobule parallel with the stem axis, and reaching more than halfway across the width of the lobe.

R. marginata, helmsiana.

Section Densifoliae Castle.

Leaves very crowded, lobules overlapping portion of the lobule of the alternate leaf above.

R. allisonii.

Section Acutifoliae St.

Leaf apices spinous or cuspidate, margins entire or toothed.

R. dentata.

In the New Zealand species there is no hard and fast line of distinction between the sections Complanatae and Adnatae.

Key to Species.

1. Microphyllous branches present (rarely absent in R. physoloba) 2.
Microphyllous branches absent 3.
2. Microphyllous branches irregular, leaves yellowish-green, plants sterile (in New Zealand) physoloba
Microphyllous branches regularly pinnate, showing on ventral side of stem, leaves glossy brown, small ribbed perianths usually present uvifera
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3. Plants with marginal gemmae, sexual reproductive organs absent (as far as is known) 4.
Plants without marginal gemmae, bearing perianths and androecia 5.
4. Dull green, leaves orbicular-oblong, somewhat falcate, apices much incurved, lobules much inflated, free angle much rounded grandis
Pale green, leaves sub-orbicular, lobules appressed, free angle somewhat elongated papulosa
5. Plants small, stems simple or sub-simple, tufted, (rare) sainsburiana
Plants medium to robust, stems irregularly pinnately to bi-pinnately branched, creeping 6.
6. Yellow-green, leaves crowded, lobule bases overlapping alternate bases allisonii
Not or rarely yellow-green, leaves less crowded, lobule bases never overlapping alternate bases 7.
7. Lobules elongated parallel with the stem axis, and reaching more than half-way across the width of the lobe 8.
Lobules not elongated parallel with the stem axis, and reaching less than half-way across the width of the lobe 9.
8. Dark-green, leaves sub-orbicular to orbicular-oblong, with a marginal rim, perianth elongate, not winged marginata
Greenish to light-brown, leaves oval-oblong, falcate, perianth winged helmsiana
9. Free ventral margin directed downwards, thereby causing the leaves to appear falcate 10.
Free ventral margin in a straight line with the keel (generally speaking), leaves not falcate 11.
10. Perianth plicate, lobule inflated with free margin folded inwards, cuticle of cells noticeably hyaline and convex plicata
Perianth tubular-elongate, lobule not or little inflated, with free angle conspicuous, cells smooth buccinifera
11. Leaves dentate and cuspidate, cells clear, 20–30 μ perianth 6–8 mm. dentata
Leaves entire, cells opaque, ca. 20 μ, perianth less than 4 mm. 12.
12. Stems to 3 cm., leaves brownish-green, approximate or contiguous, flattish, perianth gradually widening from a narrow neck levieri
Stems shorter, leaves dark-green, fleshy, imbricate, apices incurved, perianth oblong from a short neck silvosa

Radula physoloba Mont.

Rad. physoloba Mont., Voy. au Pôle Sud, 256, 1843; G. L. et N., Syn. Hep. 254, 1844; Mitt. Fl. Nov. Zel., ii, 154; Fl. Tas., 1860; Handb. N.Z. Fl., ii, 530; St., Spec. Hep., 4, 213, 1910; Rod., Tas. Bry., ii, 81, 1916; Castle in Ann. Bry., 9, 46, 1936. Rad. xanthochroma Col., Trans. N.Z. Inst., 21, 70, 1888. Rad. Colensoi, St., Spec. Hep., 4, 215, 1910.

Plants apparently sterile in New Zealand, olive- to pale yellow-green, usually on bark, may be gemmiferous. Stems 1–2 cm., irregularly pinnately branched, branches very variable in size, microphyllous amenta usually present. Cauline leaves obovate-rotund, to about 1.5 mm., convex, dorsal base attached to the stem for less than half its length, slightly auricled, then curved and extending mostly beyond the stem; apex incurved, broadly rounded, free ventral margin short, recurved, the fold being continuous with the carina; lobule large oblong-ovate, carinal portion much inflated; the base attached near the edge of the stem, for about half its length, then curved away from it; the free lateral portion appressed, the free angle obtuse; apex

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short, fairly straight; carina arched, oblique. Cells ca. 25 μ, marginal smaller, basal larger. “Perianth 2 mm. in length and 0.8 in width, the mouth slightly contracted, the lips with a conspicuous median cleft.”

I have seen no ♂ plants that could be definitely ascribed to this species.

This species is remarkable for the varied forms of microphylly which it exhibits. Plants growing on twigs and manuka may consist almost entirely of diminutive or medium-sized leaves, which may be crowded or spaced, but usually with a stem or two of ordinary leaves which help in its identification. Microphyllous branches similar to those of R. uvifera may be present on the stem, or more curious still, there may be, and usually are, terminal, capitate amenta which consist of dense clusters of very short stems, tightly packed with minute bi-lobed leaves. These may be plentiful or few, on a plant, or almost entirely absent; 2, or even all 3 of these manifestations of microphylly may be present in the same gathering.

In some instances gemmae are present. These are sub-spherical, multicellular bodies and appear to be scattered indiscriminately on the stem and surfaces of the leaves. No previous mention of these gemmae appears to have been made.

Dr. Castle has a lengthy note on R. physoloba. He says: … “The cladogynous habit, as well as other important diagnostic characters, have been overlooked, with the result that the conception of the species has been extended in general to include 2 other forms which occur within the same limits of distribution. One of these is Radula tasmanica, which was described in the Species Hepaticarum by F. Stephani, in 1910. This species is always smaller, and the carinal portion of the leaves is only narrowly inflated. It is, moreover, acrogynous and must be regarded as totally unrelated to the Montagne species. Much of the material in the Stephani Herbarium, which is included under R. physoloba must be referred to R. tasmanica. The other species which has been confused with R. physoloba is as yet undescribed. It is, however, of particular interest, since it includes the collections made by J. D. Hooker in Auckland.” (Auckland Islands.) … Dr. Castle's description of this new species will be awaited with interest, especially as R. uvifera Tayl. was also brought from the Auckland Islands by Hooker.

Assuming that Rodway's identification of R. tasmanica is correct, the following note from his Tasmanian Bryophytes ii, is significant. “This is not a distinct species, it is the form assumed by R. aneurysmalis Tayl., under favourable conditions. I have specimens in which the ends of the shoots of typical R. aneurysmalis have grown into the typical form of R. tasmanica.”

Stephani's reduction of R. xanthochroma Col., to this species is accepted by Castle, and it is very probable that R. lycopodioides Col., Trans. N.Z. Inst., 21, 71, 1888, belongs here also.

Specimens examined include 11 from the Atiamuri, Rotorua and Taupo districts, coll. K. W. Allison, 9 on manuka, 1 on tree trunk, and 1 on twigs of Dracophyllum on open plain; others are, 5 from the Wairoa district, on bark in bush, E. A. Hodgson; bark of young

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beech, Mt. Ruapehu, G. O. K. Sainsbury; Whakapapa Gorge, L. B. Moore; al318, Norsewood, Herb. Colenso; 2 with Frullania, on bark, Tauherenikau, V. D. Zotov; Mt. Arthur, G. O. K. Sainsbury; 2 from Arthur's Pass, one growing on Plagiochila circinalis, the other mixed with Dicnemon and Macromitrium sps., H. M. Hodgson; on bark with Frullania sps., Punch Bowl, Arthur's Pass, F. MacDonald; associated with mosses on beech tree trunks in forest, Avalanche Peak, Arthur's Pass, W. Martin; Silver Peak, Dunedin, G. Simpson; also from Valley of Waipoua River, North Auckland, on manuka, K. W. Allison, and Bledisloe Park, Palmerston North, E. A. Hodgson.

The type was collected in the Auckland Islands by Hombron.

Radula uvifera (Tayl.) G. L. et N.

Jungermannia uvifera, Tayl., in Lond. Journ. of Bot., 392, 1844. Rad. uvifera G. L. et N., Syn. Hep., 258, 1844; H. f. & T., Fl. Ant., 162, 1847; Mitt., Fl. Nov. Zel., ii, 154, 1855; Hanb. N.Z. Fl., ii, 530, 1807; St., Spec. Hep., 4, 231, 1910; Rod., Tas. Bry., ii, 82, 1916. Radula multicarinata G. L. et N. Syn. Hep., 258, 1844.

Plants stout, usually golden brown, creeping on bark. Stems 1–2 cm., much or little branched, amenta with small bi-lobed leaves at regular intervals on ventral side of both stem and branches, proceeding from below the lobules and usually not extending beyond the leaves. Leaves imbricate, broadly obovate-rotund, about 1.5 mm., convex when moist, with somewhat crinkly depressions when dry; dorsal base straight then curved, reaching across, to well beyond the stem, insertion very short: apices recurved, broadly rounded; free ventral margin short, a little involute, the fold being continuous with the keel; lobule ovate-oblong, base attached to near the edge of the stem for most of its length, carinal portion much inflated, the free lateral portion appressed to the dorsal lobe, free angle obtuse, apex truncate or rounded-truncate, about half the middle width; keel oblique, much arched; leaves of amenta very small, almost equally bilobed, contiguous or sub-remote. Cells ca. 20–25 μ, marginal row minute, cuticle often papillate, trigones large. Invol. leaves large, upper part rotund; lobule almost half as long as the leaf and enveloping the neck of the perianth. Perianth terminal small, subtended by innovations, easily overlooked, obovate-oblong, middle portion with numerous ridges on both faces, upper portion deflexed or decurved. ♂ bracts closely imbricate, in pairs, usually about 7, sometimes as many as 10. on ventral branches 0.9 mm. wide and quite distinct from the microphyllous amenta also present.

In the absence of perianths this species can easily be confused with R. physoloba. The golden brown colour is a good guide and the leaves, branches and amenta are very uniform in size and habit, thereby giving the plant a neat appearance. In R. physoloba, the branches may appear in all stages from microphyllous to ordinary ones, with corresponding differences in the size of the leaves.

Specimens are from: Mt. Moehau, epiphytic on kauri, 4′ up trunk, also summit of Te Moehau, L. B. Moore; Bush near Opepe, near Taupo, on tree-trunks and firewood, K. W. Allison; tree-trunks, bush on Otanepu Ridge, ca. 2,200 ft., two gatherings; on larch trunk,

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Ngapuna, near Rotorua; on tree-trunks, Roto-a-kui Bush, and edge of Pukerimu Bush, east of Taupo, ca. 2,500 ft., all coll. K. W. Allison; Waiopehu Ridge, Tararua Mts., near Levin, G. O. K. Sainsbury; also in Herb. Colenso, No. 2131, and another unnumbered.

The Handbook also gives mountainous districts in North Island, common, Colenso. Lord Auckland Islands, J. D. Hooker (type).

Also Tasmania.

Radula grandis St.

Rad. grandis St., Journ. of Linn. Soc., 29, 271, pl. 28, 1892; Spec. Hep., 4, 190, 1910; Castle, Ann. Bry., 9, 1936, issued 1937.

Plants medium, dull green, sterile, gemmiferous. Stems ca. 3 cm., pinnate to bi-pinnate but not densely so. Leaves ca. 1.5 mm., the lower ones often smaller, imbricate, ox-bicular-oblong, somewhat falcate; dorsal base reaching well across the stem; apices much recurved, either dry or moistened, somewhat revolute; marginal discoid gemmae plentiful or sparse; free ventral margin adjoining the earina usually directed downwards for a short distance; lobule very large, rhomboid, lower portion much inflated, often with a small mamillate outgrowth, free basal portion curved and more or less overlapping the stem, sometimes extending well across it, the free lateral margin curved, often touching or over-reaching the carina of the lobule directly above it; angle obtuse; earina well-arched, oblique. Cells ca. 20 μ, apical marginal minute. Perianth unknown.

This is a fine plant. Stephani notices that it resembles small forms of Porella, stangeri, but neither in the Linnean Journal or the Species Hepaticarum does he mention the marginal gemmae. These, together with the much recurved leaf-tips, and the decurved free ventral margin distinguish it from R. physoloba and R. uvifera. The dull green colour, too, is constant and characteristic, except that one specimen from Morere Bush is a brownish-green and with scarcely any gemmae.

Localities are: Mt. Ruapehu, G. O. K. Sainsbury; on earth, dry, shady bank, east of Taupo, ca. 2,500 ft., K. W. Allison; on manuka, near Atiamuri, K. W. Allison; tree trunks in Puaiti Bush, near Rotorua, and Rxiahakune Bush, near Taupo, ca. 2,500 ft., K. W. Allison; on trunk of large tree, bush by Maungapoike Falls, near Wairoa, E. A. Hodgson; Waiopehu Ridge, Tararua Mts., G. O. K. Sainsbury; Wilton's Bush, near Wellington, R. Mason; three gatherings from Marlborough, localities not stated, J. H. McMahon; also from Bush near Queen Charlotte Sound, J. H. McMahon.

Stephani gives New Zealand, North Island. Castle speaks of it as “of Auckland and New Zealand.” I have no knowledge of R. grandis having been collected in the Auckland Islands. Perhaps the name has been confused with the Auckland of the North Island of New Zealand.

Radula papulosa St.

Rad. papulosa, St., in Journ. of Linn. Soc., 29, 272, 1892; Spec. Hep., 4, 189, 1910. Rad. complanata. Dum. in Handb. N. S. Fl., 2, 530, 1867.

Plants pale olive- to pale yellow-green, creeping in mats on bark (one specimen creeping-over Papillaria crocea attached to the bark). Stems 1–2 cm., fairly closely pinnately branched, longer branches

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again bearing pinnules. Leaves slightly to closely imbricate, sub-circular, slightly convex, margins somewhat recurved or plane, occasionally hyaline, often crenulate and gemmiferous, being edged with irregularly shaped, multicellular bodies, originating from the marginal cells; the free dorsal part extending across the stem and overlapping the basal portion of the opposite leaf; the free portion of the ventral margin more or less in a straight line with the keel. Lobule large, rectangular-rhomboid, somewhat oblique, scarcely arched, base straight, attached to the stem axis for about half its length, the free portion overlapping the stem, but rarely reaching half way across it; apex truncate, free angle obtuse, often somewhat elongated; a small, brownish, mammilate outgrowth sometimes present on the mid-carinal portion. Cells rounded-hexagonal, rather obscure, ca. 20–25 μ; no trigones. Fructification unknown. Androecia in 3–4 pairs on small branches, may be basal or terminal, bracts about half the size of the ordinary leaves or less.

The sub-circular imbricate leaves with gemmiparous margins make this plant easy of recognition.

As Stephani points out in the Linnean Journal, his R. papulosa is most probably the plant described in the Handbook as R. complanata, a gemmiferous plant of the Northern Hemisphere. R. complanata is not the same species as R. papulosa, as is shown by English specimens in my possession, and it is not likely that both species occur in New Zealand.

My specimens are all local: Base of cabbage tree (Cordyline australis), Alton, near Wairoa. In bush Te Tiki, ca. 1,500 ft., inland from Wairoa; in bush, Terapatiki, near L. Waikaremoana, two gatherings, one ♂ in bush by Maungapoike Stream, near Wairoa; also in Herb. Colenso, No. a1643*.

The Handbook plant is from Ruakawa, H. B., coll. Colenso. Stephani gives no definite locality for his type.

Radula sainsburiana Hodgson and Allison sp. nov.

Paroica, minima, caespitosa flavo-rufescens. Caulis ad 1.3 cm., longus, simplex vel parum ramosus. Folia caulina ovato-elliptica, parum oblique-patula, concava; basi ampliato, caulem bene tegente et supra extento; lobulus late ovatus, inflatus deinde appressus vel involutus, apice angusto, excurrente in marginem incurvum folii. Folia ramea minora, erecto-patentia, valde concava; carina bene arcuata, sinuata. Cellulae ca. 20 μ, parietibus incrassatis. Perianthia innovata, oblonga 1.5 mm. 0.1 mm., ore truncato. Androecia infraperianthium, bracteis 3–4 jugis.

Plants paroicous, very small, pale yellow-green to reddish-brown with tints of rose, tufted, with Sphenolobus perigonialis (also rosetinted). Stems to about 1.3 cm., simple or little branched. Leaves concave, ca. 0.6 mm., loosely imbricated, spreading, similar in appearance to those of Lejeunia; dorsal lobe oval to ovate-rotund, base arched, extending beyond the stem and overlapping a small portion of the alternate lobe above, the free ventral margin directed upwards, slightly incurved, the fold continuous with the carina; lobule more

[Footnote] * A further locality is Bledisloe Park, Palmerston Nth., E. A. H., Oct., 1944.

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or less broadly ovate, base seems to be attached to the stem for most of its length, line of attachment at varying distances from the margin of the stem, carinal portion inflated, the free side appressed; apex narrowed and usually continuous with the slightly incurved ventral margin of the lobe, no rhizoidal outgrowths. Carina strongly arched, sinuate. Cells ca. 20 μ, walls thick, cuticle papillate distinctly showing along the edge of the carina; trigones confluent. Branch leaves more erect, lobes may be much smaller, with the appearance of being laterally extended; lobules remaining much the same size. Invol. leaves larger than the foliage, lobule longly obovate, extending half way or more across the width of the lobe. Perianth oblong, ca. 1.5 mm. × ca. 0.8 mm., subtended by an innovation and 3 or 4 perigonial bracts with closely imbricated lobules; mouth two-lipped, truncate. (Two inflorescences seen.)

This little species is distinct in its small size, pale colour, spreading leaves and paroicous inflorescence.

The habitat seems to have been on rock; locality, creek-bed, Mt. Ruapehu, coll. G. O. K. Sainsbury.

A duplicate of the type is in the Herbarium of the Plant Research Bureau, Wellington.

Radula allisonii Castle.

Rad. Alstonii Castle in Hep. Select. et Crit., in Ann. Bry., 8, 1935, corrected to Rad. Allisonii in Ann. Bry., 10, 130, 1937.

Plants medium to robust, pale yellow-green, densely creeping, often mixed with R. buccinifera, usually on bark. Stems ca. 1.5–2.5 cm., irregularly pinnately ramose, branches of varying length. Leaves large on the main stem, ca. 1–1.2 mm., medium to small on the branches, imbricate; dorsal lobe concave, oval, with the rounded apical portion irregularly recurved or even revolute (when dry), base from a straight insertion, reaching beyond the stem about 1.5 mm., and overlapping the base of the alternate leaf above; the free ventral margin rarely directed downwards, in the case of the smaller branches, may be directed upwards. Lobule outstanding in both size and shape, not inflated; base attached to the stem for about half way in a more or less straight line, then expanding broadly in a curve which mostly overlaps the basal portion of the alternate lobule above, and is continued, to form the free lateral margin of the lobule; the free angle usually obtuse, may be acute or acuminate; apex lunate or sinuous, rarely truncate; carina oblique, fairly straight; tufted rhizoidal outgrowths may be present. Cells ca. 22 μ, cuticle papillate, trigones minute or absent. Invol. leaves sub-erect, lobule reaching about half the length of the lobe. Perianth oblong, ca. 3.5–4 mm., × ca. 0.9–1 mm., margins partially or entirely irregularly winged, wings to 0.1 mm., apex often decurved, mouth truncate, straight to sub-crenate; subtended by innovations. ♂ bracts small to medium, on branches of varying lengths, both pinnae and pinnulae.

The pale colour, crowded leaves, with large lobules deflected from the stem axis, and winged perianths, are the distinguishing features of this notable species, of which the distribution as far as is known, is restricted to the Central North Island Plateau District. Of the 15 specimens in Mr. Allison's herbarium, 7 are from tree trunks, Puaiti

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Bush, South of Rotorua; 2 from tree trunks Pukerimu Bush, ca. 2,500 ft., East of Taupo; 2 from rock, East of Waiotapu Valley, ca. 1,600 ft., Rotorua Co.; tree trunk, Roto-a-Kui Bush, East of Taupo; tree trunk, Ruahakune Bush, near Taupo; on earth, under manuka (Leptospermum scoparium), near Atiamuri; on bark, bush at edge of Urewera; also on bark in bush, Taihape Domain, E. A. Hodgson.

Radula marginata (Tayl.) G. L. et N.

Jungermannia marginata Tayl., in Lond. Journ. of Bot., 566, 1844. Rad. marginata G. L, et N., Syn. Hep., 261, 1844; Mitt., Fl. Nov. Zel., ii, 154, 1855; Handb. N.Z. Fl, ii, 530; St., Spec. Hep., 4, 230, 1910.

Plants dark green, rarely fruiting, in depressed mats on rock, earth, bark or logs. Stems to about 3 cm., little or much branched. Leaves orbicular-oblong, imbricate, slightly convex, margins having a thickened, somewhat horny appearance, due to the differentiated cells; base of dorsal lobe about half attached to the stem, then rounded and extending most of the way across it; free ventral margin in a straight line with the carina, which is short, though somewhat longer in the Marlborough specimen; lobule ovate-elongate, parallel with the stem, angle obtuse, base about a quarter attached to the stem; tufted rhizoids may be present. Cells obscure, marginal minute and crowded, median ca. 20–25μ, trigones small or absent. Invol. leaves more oblong than the cauline, with an even larger lobule. Perianth ca. 4.5 mm., including the neck, 0.9 mm. wide, lips truncate. ♂ bracts small, in 4–5 pairs, on very short branches, or at the bases of longer ones.

The only fruiting plants I have seen are from Waihau Bay or Raukokore River, Bay of Plenty, coll. G. M. O'Malley, No. H 638 Herb. Allison; bare rock in stream, Waipoua Forest, K. W. Allison.

The large, rounded, deep-green leaves with a noticeable rim, together with the unusually tall lobule, enable this species to be easily recognised.

Other localities are: Common on rock in shaded wooded gullies, Mt. Wellington lava field, Auckland, E. B. Aschroft, comm. K. W. Allison; Titirangi, Auckland, E. D. Swanberg; Shortland, Thames, T. Kirk, 1868, No. 6071, P. R. B. Herb.; on log between wet cliffs, bush at Maungapoike Falls, Wairoa; on earth mound, Morere Bush, E. A. Hodgson; upper Tiritea River, V. D. Zotov, No. 6889 P. R. B. Herb.; wet rock, Wilton's Bush, Wellington, R. Mason; Kapiti Island, with Plagiochila stephensoniana, No. 4077 P. R. B. Herb., A. H. Wilkinson; Red Hills, Marlborough, ♂, collector uncertain.

The type was from the Bay of Islands, coll. Hooker. The Handbook also records it from Nelson, coll. Mantell.

Radula helmsiana St.

Rad. Helmsiana St., 4, 231, 1910.

Plants medium, brownish-green when fresh, usually light brown when dry, creeping on bark. Stem 1–1.5 cm., little or much bipinnately branched, crowded or extended. Leaves oval-oblong, inclined to be falcate, contiguous or more usually a little imbricate, may be flat, when the plant greatly resembles Lopholejeunia plicatiscypha, or bent or curved, generally upwards; ca. 0.9–1.1 mm.; × ca. 0.7 mm.; dorsal base sub-truncate, less than half attached to the stem, extending rarely less than half-way across, but not beyond it;

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free ventral margin in a straight line with the carina, or deflected somewhat, from a wide sinus; lobule large, rhomboid-ovate, often reaching beyond the carina of the lobule above it; base long, attached to the stem for less than half its length; the free angle obtuse; carina oblique, only slightly arched. Cells, marginal ca. 12 μ, median ca. 20 μ, with a dotted appearance; trigones minute. Invol. leaves oblong, may be twice as long as broad, lobule reaching about three-quarters across the width of the lobe. Perianth ca. 2.4 mm. long, oblonglanceolate, margins winged, usually widest below the middle, ca. 0.8 mm. excluding the wings; wings ca. 0.1 mm. or less, may be undulate; mouth ca. 0.5 mm. wide, two-lipped, lips with a median cleft. ♂ stems all lying in the same direction, bracts to 30 or more pairs, though few appear to contain antheridia; lobe erect, the upper part spreading, lobule almost as large as the lobe.

There is an element of doubt about this identification, as the leaves of Stephani's plant, which unfortunately was a sterile one, are described as “subrotunda”; but the large, elongate lobule is the same. It is just possible that Stephani has mistaken a discoloured specimen of R. marginata for a new species. If there really are 2 separate species involved, then a new name must be found for the North Island plant (described above).

Distinct from R. allisonii in the differently shaped lobule, the less crowded, oval-falcate leaves; and the perianths though winged, are not truly oblong as in that species, but narrowed towards the base and apex, while the lips are cleft in the middle.

Localities are not numerous. Common in bush at Morere Hot Springs, between Wairoa and Gisborne; “McKinnon's” Bush, bush at Maungapoike Falls, both near Wairoa, E. A. Hodgson; forest between Lake Rotoehu and coast, K. W. Allison; Puaiti Bush, near Rotorua, K. W. Allison; Wilton's Bush, Wellington, 2 gatherings, R. Mason; Bledisloe Park, Palmerston North, on bark, E. A. Hodgson.

Radula plicata Mitt.

Rad. plicata Mitt., F1. Nov. Zel., ii, 154; Handb. N.Z. F1 ii, 530, 1867; St., Spec. Hep., 4, 213, 1910; Pears., Univ. Cal. Pub, Bot., 10, 1923.

Plants dull green, faintly brownish, closely matted. Stems 2–3 cm., pinnately to bi-pinnately branched. Leaves 0.8–1 mm., imbricate, convex, somewhat falcate; dorsal lobe with a somewhat truncate base produced into an ample curve which may considerably over-reach the stem; apex recurved; free portion of the ventral margin extending downwards from the oblique lobule at approximately a right angle; lobule medium, lower portion inflated, longly decurrent, sub-rhomboid, base adnate with the stem for practically its whole length, the free latteral marginal area usually folded inwards over the inflation, which may be narrowed towards the rounded-truncate apex; the free angle obtuse; earina strongly arched or sinuate, oblique. Cells ca. 20 μ, larger at the base, trigones very small; cuticle convex and hyaline, showing along the edge of the carina and the backward fold of the upper part of the dorsal lobe. Invol. leaves somewhat smaller than the cauline, obovate, dorsal lobe not produced downwards, lobule shorter and broader than the cauline, flat. Perianth ca. 3.5 mm., oblong, gradually widening from a narrow neck, with three full-

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length plicae on the dorsal and 2–4 on the ventral face. Androecia in 4–6 pairs on short branches, or intercalary on longer ones. Stephani speaks of the branches as srery long, sometimes occupying the whole plant.

The plicate perianth will always distinguish this species which, as Mitten points out, very closely resembles R. buccinifera. Another difference is that the lobule in R. plicata is longer, that is, the free angle is further removed from the stem, and the carinal portion is more inflated. The leaves are less, but more regularly recurved, and lack a certain lustre natural to those of R. buccinifera.

I certainly agree with Pearson, who wrote about Setchell's plant from Mamaku, that Stephani's note about the perianth folds being not very noticeable, and only on the older branches, is incorrect.

Fruiting plants seen are from Karamea, Nelson, coll. Miss Foot, and Dreamlands, Waitakere Ranges, Auckland, L. B. Moore, No. 24763, P. R. B. Herb. A ♂ specimen, epiphyllous on Trichomanes elongatum is also from Dreamlands, No. 24729, P. R. B. Herb., coll. L. B. Moore. A sterile plant enveloping stems of Weymouthia cochleanfolia (moss), is from near Fox Glacier, coll. Mrs. Knight; while 2 sterile plants which I think belong here, but cannot say with certainty, are from Wilton's Bush, Wellington, R. Mason, and Little Barrier Island on Hypopterygium filiculaeforme, also a moss, coll. W. M. Hamilton.

The type locality is given as Auckland, coll. Dr. Sinclair. Radula buccinifera (Tayl.) G. L. et N.

Jungermannia buccinifera Tayl., Hep. Antarct., in Lond. Journ. of Bot., 580, 1844. Rad. buccinifera G. L. et N., Syn. Hep., 261; Mitt., F1. Nov. Zel., ii, 154; F1. Tas., 1800; Handb. N.Z. F1, ii, 530; Rod., Tas. Bry., ii, 81, 1916; St., Spec. Hep., 4, 227, 1910.

Plants dioicous, in compact, spreading mats when creeping on bark or rock, glossy golden brown to olive green, sometimes gemmiferous. Stems 2–3 cm., freely branched, branches and branchlets short or long, leaves on the short ones may be smaller and more erect, but not minute. Cauline leaves convex, ovate-falcate, less than 1 mm., antical base more or less truncate, then curved, often extending beyond the stem; apices irregularly recurved, giving the leaves a collapsed look; leaf edge may be sinuate or more rarely still, with a blunt serration or other marginal irregularity; free ventral margin directed downwards, forming a sinus with the lobule; lobule medium or small, vertically rhomboid, base slightly inclined towards the middle of the stem, long, attached to the stem for most of its length, straight or a little curved at the top; the free apical angle, obtuse or bluntly acute, may be somewhat elongated, or curved in the direction of the stem; usually flat, may be inflated in the lower portion, or bulging, on the branches; carina oblique, more or less arched. Cells ca. 10–20 μ, marginal minute, cuticle with a hyaline, low papilla. Invol. leaves obovate, lobule with a very long base. Perianth ca. 3 mm., narrow-tubular, widened somewhat near the apex, month crenate or truncate; sterile perianths shorter and broader.

Perigonal bracts usually in 3 or 4 pairs, alternating with 3 or 4 “proper” leaves on fairly long branches, or on short, purely ♂ branches; lobule about half the size of the lobe.

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A plant from bush by Makaretu Stream, near Wairoa, growing on bark, has the lobules filled with multicellular septate gemmae. When moistened these float out into the concave surface of the leaf. Another plant, on firewood, Bay of Plenty, coll. K. W. Allison, is ♂, and has gemmae in the lobules of the “proper” leaves alternating with the perigonal bracts. These are shorter than those of the Wairoa plant. A third plant, ♂, from near Atiamuri, on earth under manuka, has but a few gemmae in some of the lobules.

Specimens examined include 23 from Taupo, Rotorua and Atiamuri districts, habitats being bark of tree-trunks and branches, treeferns, rocks, on logs, on earth under manuka, one on Mt. Ngongataha creeping over fronds of Hymenophyllum demissum, also one on bare bark of rimu, 6–8 feet from ground, between Lake Roto-ehu and coast, Bay of Plenty, all coll. K. W. Allison. Other localities are: Bush by Lakes Waikaremoana and Waikare-iti, on bark and Hymenophyllum sp. respectively; Morere Bush; Maungapoike Falls Bush, enveloping Polytrichum juniperum, all coll. E. A. Hodgson; Mt. Maungapohatu, B. Teague; Mangahao Headworks on Cyrtopussetosus, G. O. K. Sainsbury; headwaters of east branch of Hawdon River, North Canterbury, H. M. Hodgson; with Plagiochila lyallii near Fox Glacier, Mrs. Knight.

The Handbook also gives Otago, Hector and Buchanan.

The type was collected in Tasmania by Hooker. Also found in Australia.

Radula dentata Mitt.

Lejeunia dentata Mitt. in F1. Nov. Zel., ii, 159, 1855. Radula dentata Mitt., M. S.,; Handb. N.Z. F1., ii, 1867; St., Spec. Hep., 4, 156, 1910.

Plants medium, terrestrial, glossy, light brown or olive green. Stems to about 3 cm., rather closely pinnately branched. Leaves imbricate, convex, varying in size, usually less than 1 mm.; dorsal margin 1–3 spinous-toothed, or dentate or sinuous, the free basal portion reaching from about one-third to half the distance across the stem; apex recurved with 1–3 spinous teeth; free ventral marginal portion entire or toothed, not directed downwards from the lobule; lobule ovate, oblique, carinal portion much arched and inflated, the free margin appressed to the lobe, the free angle usually obtuse. Cells ca. 20–30 μ, no trigones. Invol. leaves in 3 pairs, the upper-most much larger than the cauline, the lower pairs smaller, entire or cuspidate or with few teeth. Perianth very long, 6–8 mm., from a narrow tubular base, middle width ca. 0.5 mm., mouth slightly wider, truncate.

I cannot help but think that R. cuspidata St., Spec. Hep., 4, 156, also from the Great Barrier Island, is only a form of R. dentata. The only difference of any possible consequence that I can infer from Stephani's description and drawing, is that R. cuspidata lacks the marginal teeth; the shape of the leaf and lobule and the copious branching are apparently the same, while the perianth is not mentioned at all; but in some of the lower leaves of the Moehau plant the marginal teeth are almost obsolete, and Stephani himself says under R. dentata, “Die Blattzähne variinen ausserordentlich in Anzahl

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wie in Grosse und es kommen sogar völlig gansrandige Blätter vor.” Stephani's leaf-outline, with its longly cuspidate incurved apex is strikingly similar to leaves of R. dentata.

A well-marked species. Localities: Great Barrier Island (in Kirk's collection), mixed with Blepharostoma sp., and Chiloscyphus triacanthus; Te Moehau Mt., ca. 2,000 ft., also mixed with C. triacanthus, L. B. Moore; growing in short, thick clump of Plagiochila. on sandy creek bank, Waipoua Forest, K. W. Allison.

Stephani gives New Zealand and New Caledonia. The type locality is given as Auckland, coll. Sinclair.

Radula levieri St.

R. Levieri St. Spec. Hep., 4, 227, 1910.

Plante usually dioicous, olive-green to light brown, straggling or compact on bark, rock, earth, or growing on, or mingled with, mosses or other hepatics. Stems ca. 3 cm., thin, reddish-brown or darker, branched pinnately, branches of varying lengths. Leaves 0.9–1 mm., usually flat or almost so, little altered when dry, elliptic or elliptic-orbicular, approximate, contiguous or less often, slightly imbricate; dorsal line of attachment straight, free portion curved a little but not extending to the middle of the stem; free ventral margin in a straight line with the carina, not proceeding downwards; lobule small, sub-quadrate, base long, mostly joined to the stem, angle obtuse or sub-acute; carina oblique, straight or slightly inwardly curved, longly decurrent; branch leaves usually smaller; cells 6-sided, opaque, ca. 20 μ, trigones absent or very small. Invol. leaves smaller than the cauline, oval, lobule reaching almost across the width of the leaf. Perianth to 3.5 mm., gradually widening from a narrow-tubular neck, or oblong from a less elongated neck, mouth truncate or crenate. ♂ bracts in a few pairs on short pinnate branches, or numerous on long ones, imbricate, more erect than the ordinary leaves, lobules imbricate reaching half to two-thirds up the lobe.

A plant from clay bank, Otari Reserve, Wellington, coll. N. Kemble Welch, comm. K. W. Allison, is definitely monoicous, but this does not appear to be usual.

I think it is very probable that R. strangulata Tayl. in Land. Journ. of Bot., 1846, Syn. Hep., 730, but omitted from the Flora Novae Zelandiae and the Handbook, is the ♂ plant of R. levieri St. Colenso draws attention to this species in Trans. N.Z. Inst., 21, p. 71. It was collected at Bay of Islands by Hooker.

R. levieri can be distinguished from R. buccinifera by the flattish non-imbricate, non-falcate leaves, whose dorsal bases do not reach across the stem, which thus appears as a wavy line between two rows of alternate leaves.

North Island localities are numerous, extending from North Auckland to Wellington. South Island stations are as follows: Bay of Many Coves, Marlborough; Banks Peninsula, both coll. H. M. Hodgson; watershed of Kaituna Creek, Nelson, G. Simpson and J. S. Thomson; water-race in Silverstream Valley, also Whare Flat, Dunedin, G. Simpson and J. S. Thomson; near Dunedin, M. Finlayson.

Stephani's type was from Westland, coll. Helms; also collected by Beckett.

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Radula silvosa Hodgson and Allison sp. nov.

Dioica fusco-virida, implicata. Caulis ca. 1 cm., irregulariter pinnatus vel bi-pinnatus. Folia caulina imbricata, carnosa, elliptica, ca. 0.8 mm. longa, vel late sic; apico incurvo, basi breviter accreta, supero libero, ampliato, caulem tegente; lobulus sub-quadratus basi ad marginem caulinis longe accretus, supra basin interdum leviter ampliatus; angulo obtuso apice oblique truncato; carina obliqua, substricta longe decurrente. Cellulae ca. 20 μ, obscurae. Perianthia utrinque innovata, oblonga, 1.5–2.5 mm. × 0.8–1.1 mm.; ore truncato vel crenato. ♂ rami breves, pinnati remotiuscule, bracteis 3–4 jugis.

Plants in layered mats on earth mounds and overground roots usually in deep shade, often in association with the mosses Homalia pulchella and Eriopus brownii, dark green, fleshy when fresh. Stem ca. 1 cm., irregularly pinnately to bi-pinnately branched, sub-floral innovations of varying lengths. Leaves imbricate, fleshy, oval to ovalrotundate, apices may be incurved; dorsal base shortly attached, then arched and usually reaching more than half-way, sometimes wholly, across the stem, and often covering a portion of the alternate lobe above it; free ventral margin not, or rarely, directed downwards, usually in a direct line with, or continuing the curve of, the carina; lobule not inflated, triangular-rhomboid, base long, sometimes reaching 3.5 mm. in length, the upper, free part sometimes slightly overlapping the stem; the free angle obtuse or sub-acute, the apex obliquely truncate, always shorter than the base and never quite parallel with it; carina oblique, scarcely arched, longly decurrent. Cells hexagonal, ca. 20 p, the lumen partly or wholly filled with dark, opaque chloro-plasts. Trigones O. Invol. bracts larger, lobule elongate, reaching about three-quarters of the way across the width of the lobe. Perianth 1.5–2.5 mm. × 0 8–1.1 mm., occasionally somewhat widened towards the mouth, flat, with traces of rudimentary wings; mouth truncate or crenate. (In a specimen from Morere bxish, the perianths are funnel-shaped with crinkled mouths.) ♂ bracts in 3 or 4 pairs on short pinnate branches.

Nearest to R. levieri, from which it is sometimes difficult to distinguish in the dry state, the shape and structure of the lobe and lobule being similar in both species. But the leaves of R. silvosa are more crowded, with the dorsal lobe overlapping the stem, and the whole plant is stouter and more compact. The perianth, too, is shorter and broader, without the elongated tubular neck of R. levieri, though occasionally the perianths do approach those of the latter species in shape. When freshly gathered, the dark green, fleshy leaves are a great help in identifying the plant.

Not uncommon in bush in Wairoa County. The type is from Te Tiki Station, Wairoa, ca. 1,500 ft., No. 24766 P. B. B. Herb. Also from Ohope, near Whakatane, H. M. Hodgson, and Wilton's Bush, Wellington, B. Mason.


In Species Hepaticarum 4, 190, Stephani ascribes the Norfolk Island species, Radula mittenii St., to New Zealand. I have not yet met with any plant that corresponds with his description and drawing. According to these, the plantmay be 5 cm. long, the leaf re-

Picture icon

Fig. 1—R. allisonii. Figs. 2–3—R. sainsburiana. Fig. 4—R. plicata. Fig. 5—R. helmsiana. Fig. 6—R. buccinifera. Fig. 7—R. papulosa. Fig. 8—R. physoloba. Fig. 9—R. marginata. Fig. 10—R. uvifera. Fig. 11—R. silvosa. Fig. 12—R. grandis. Fig. 13—R. levieri. Fig. 14—R. dentata. blank

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sembling that of R. levieri, and the perianth that of R. buccinifera, but it differs from both these species in the lobule which has the free portion of the lobule base ampliate, and partly covering the stem. A specimen in Herb. Petrie, No. 131, determined by Stephani as R. mittenii, is R. levieri.

Radula albipes Col., Trans. N.Z. Inst., 21, p. 71, 1888, is Lejeunia nudipes. Tayl, according to Stepheni in his Revision of Colenso's New Zealand Hepaticae, and from the description, this appears to be correct.


In concluding I gratefully acknowledge the loan of Mr. K. W. Allison's entire collection of Radulas, and of still further consignments of liverworts from the Herbarium of the Plant Research Bureau. Thanks are also extended to Miss L. B. Moore for helpful criticism of the manuscript.

Literature Consulted.

Castle, H., 1937. A Revision of the Genus Radula, 1. Ann. Bry., 9, 13–56, Leiden.

Colenso, W., 1888. Description of Newly Discovered Cryptogamic Plants. Trans. N.Z. Inst., 21, pp. 70–72.

Gotische, C. M., Lindenberg, J. B. G., et Nees, C. G., ab Esenbeck, 1844–1847. Synopsis Hepaticarum, including Supplement, Hamburg.

Hooker, J. D., Botany of the Antarctic Voyage, London. cum MITTEN, W., 1867. Handbook of New Zealand Flora, 2, London.

Mitten, W., 1855. Flora Novae Zelandiae 2, London.

Pkabson, W. H., 1923. Notes on a Collection of New Zealand Hepaticae. Univ. Cal. Pub. Bot., 10, 317.

Rodway, L., 1916. Hepatics. Tas. Bry., 2, Hobart.

Stephani, Fr., 1910. Species Hepaticarum 4, Geneva.

—– Hand-drawings of Species (Icones), ined.

—– 1892. Colenso's New Zealand Hepaticae, Journ. Linn. Soc., 29, pp. 263–280.

Taylor, T., 1847. Flora Antarctica, London.


Tile following typographical corrections should be made to Hodgson, E. A., 1944. A Review of the N.Z. Species of Plagiochila. Trans. Roy. Soc. N.Z., vol. 73, pt. 4:—

Page 291, line 6: should read “the whole length of the stem …”

Page 291, line 17: should read “… strongly-toothed.”