Review of the New Zealand Species of Schistochila, with notes on Colenso's Species.
The statements and descriptions in this paper are based on the combined study by Mr. K. W. Allison, of the State Forest Service, and myself, over a prolonged period, of the New Zealand liverworts—research work made difficult by the absence from this country of both types and herbaria. However, the late Dr. Marshall Howe, of New York, kindly sent us fragments of Mitten's types from the Bronx Park Herbarium, and these, which are now in the Plant Research Bureau Herbarium, have been of the greatest assistance, though not in connection with this paper. Nor can overseas bryologists give much reliable assistance, as their acquaintance with New Zealand plants is usually a superficial one, though Dr. Th. Herzog, of Jena, has given much of his valuable time to the study and identification of plants and the description of new species, and Dr. Fr. Verdoorn has helped with the section Holostipae of Lejeunia, while Mr. W. E. Nicholson, of Lewes, Sussex, has consistently helped us in many ways.
As regards literature, Sir Joseph Hooker's Handbook of the New Zealand Flora, Part ii, has formed the basis of our work. We have also had at our disposal the Synopsis Hepaticarum of Gottsche, Lindenberg and Nees, published in 1844, and three volumes of Hooker's Botany of the Antarctic Voyage, viz., Flora Antarctica, Flora Novae Zealandiae, and Flora Tasmaniae, published in 1847, 1855, and 1860, respectively. I understand that these old and valuable books were procured for the library of the Hawke's Bay Philosophical Society by the late Rev. William Colenso; and it is most fortunate that they escaped the destructive earthquake fire of 1931.
All efforts to obtain a copy of Band 9 of Engler and Prantl's Pflanzenfamilien proved unsuccessful, though a second edition was supposed to be in course of preparation some years before the war. Stephani's Species Hepaticarum has also been unobtainable, but this work is really indispensible, and through the friendly interest of the late Sir Arthur Hill, Director of Kew, I was enabled to obtain manuscript copies of the descriptions of New Zealand plants, including Stephani's new species. Copies of both drawings and descriptions were also procured from Fräulein Stephani, of Leipsic, and Mr. Nicholson has also kindly copied both drawings and descriptions for us. Even so, the list is still incomplete.
The Students' Handbook of British Hepatics, by S. M. MacVicar, is particularly useful for general information, but unfortunately it has been out of print for some time. A copy of this work was obligingly lent to us by the Librarian of the Auckland Institute and Museum, per Miss Cranwell.
I would take this opportunity of acknowledging our indebtedness to Mr. G. O. K. Sainsbury, of Wairoa, who from the outset has been a tower of strength in his unfailing encouragement and advice.
A very great deal still remains to be done in the systematic study of New Zealand Hepaticae and further gatherings of plants are most welcome, specially from the South Island.
Genus Schistochila Dumortier.
The genus Schistochila, usually considered the most beautiful of the hepatic genera, includes something over 100 published species, 11 of which are shortly described in Part ii of Sir J. D. Hooker's Handbook of the New Zealand Flora, published in 1867, under the generic name of Gottschea. Dumortier's earlier name of Schistochila has now been restored and will be used throughout this paper. The old name of Jungermannia, used by Sir William Hooker in the Musci Exotici and by Dr. Taylor in the London Journal of Botany was the general name for all the Hepaticae at that time.
Plants small to robust, stems simple or with few branches, sub-woody or fleshy, with or without paraphyllia, and with rhizoids on the ventral side or at the base. Leaves small to very large, succubous, bi-lobed, complicate, the dorsal lobe smaller than, or equal to, the ventral, which is usually referred to as the leaf, and attached to the latter at varying distances from the margin, the portion of the ventral lobe below the line of insertion, thus forming a wing; sometimes lamellate, margins serrate, ciliate or sub-entire. Stipules present or absent, the armature conforming with that of the leaves. Sporophyte deeply sunk in the swollen apex of the stem, which thus forms a tubular involucre adnate with the calyptra and overlaid with leaves.
In most of the species there is marked variability in the armature of the leaf margins.
Key to Species.
Group A. Stipules present.
[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]
|1. Leaves lamellate||2|
|Leaves without lamellae||7|
|2. Leaves large, incised, so as to be pinnatifid||appendiculata|
|Leaves not, or scarcely incised, smaller||3|
|3. Stems without paraphyllia, leaves toothed||repleta|
|Stems with paraphyllia||4|
|4. Leaves pubescent||glaucescens|
|5. Lobes sub-equal, stipules 4-fid, with ciliate apices||lehmanniane|
|Dorsal lobe shorter, truncate||6|
|6. Leaves ciliate-dentate, stipules 4-fid ciliate||balfouriana|
|Leaves slightly ciliate-dentate, stipules 2-fid||unguicularis|
|7. Stems more than 3 inches, with axillary tufts of scales||nobilis|
|Stems less than 3 inches, without tufts||8|
|8. Plant thick and fleshy||splachnophylla|
|Plants with leaves not thick and fleshy||9|
|9. Dorsal lobe shorter, muricate||muricata|
|10. Leaves ciliate all round with hair-like cilia, stipules 2–4-fid, ciliate||ciliata|
|Plants small, leaves and stipules not ciliate all round||11|
|11. Leaves sub-entire||colensoana|
Group B. Stipules absent.
|1. Stems 2–3 inches high, branched, leaves pinnatifid, toothed and ciliate||pinnatifolia|
|Plants smaller, leaves non-pinnatifid, lobes concave, inflated||2|
|2. Leaves papillose, very pale green||tuloides|
|Leaves smooth, brownish||conchophylla|
Descriptions of Species.
Schistochila appendiculata (Hook.) Steph.
Jungermannia appendiculata Hook., Musci Exotici, 1818. Gottschea appendiculata G., L. et N., Syn. Hep., 1844; Mitt., Nov. Zel., ii, 1855; Handb. N.Z. Fl., ii, 1867. G. moniliformis Col., G. winkelmanii Col., Trans. N.Z. Inst., xxi, 1888. Schistochila appendiculata Steph., Spec. Hep.
Plants robust, yellow- to olive-green. Stem 5–15 cm., sub-woody, procumbent at base, then erect, simple or with 1 or 2 branches, often bare in the lower part, no paraphylls, but with red rhizoids at the base. Leaves very large, to 9 mm. long by 5 mm. broad, oblong-ovate, incised and pinnatifid towards the apex, margin serrulate, posterior basal, entire, lamellae running obliquely inwards from the incisions, dorsal lobe from ½ to ⅔ the length of ventral, semi-ovate, acute, sometimes almost bidentate, margin strongly arched from the base, overlapping the stem, undulate, serrulate towards apex, line of insertion oblique, wing definitely single. Stipules large, bifid to the base, segments ovate, margins recurved and coarsely laciniate-toothed. Cells variable in size, ca. 20–30 μ × 10–15 μ, basal ca. 70–75 μ × 25 μ. Involucre cup-shaped, the mouth cut into 8–9 equal toothed segments, the lower tubular part, to the inside walls of which the calyptra adheres, overlaid with bi-lobed leaves.
The Handbook states that the leaves of this species are without lamellae; but this is not the case as narrow toothed lamellae are certainly present.
The most handsome and striking of the species, found on the ground in forest shade from the far north to Stewart Island, also Kapiti Island, but not particularly common.
Schistochila nobilis (Hook.) Steph.
Jungermannia nobilis Hook., Musci Exotici, 1818. Gottschea nobilis Nees; G., L. et N., Syd. Hep., 1844; Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., ii, 1867. Schistochila nobilis Steph., Spec. Hep.
Plants robust, yellow-green, in small or widespread patches. Stem from 6 to 20 cm., procumbent in lower half, then erect or flexuose, almost woody, very brittle when dry, in large specimens black and bare in lower part, in smaller plants small rhizoids present for a short distance amongst old and broken leaves; leafy part with axillary tufts of toothed and ciliated paraphylls, often dichotomously branched. Leaves 4–6 mm. long, 3 mm. broad, imbricate, without lamellae, ovate, acute, serrulate-ciliate at base of ventral margin, dorsal lobe ¼ shorter, broadly semiovate, acute, strongly arched at the base and reaching well above the ventral lobe and overlapping the stem, thus giving the plant a very characteristic look from the dorsal aspect; margin undulate, line of insertion sub-parallel with the lower margin of the leaf. Cells
very variable in size, 15–30 μ, basal 60–80 × 25 μ Trigones small. Stipules 2½ mm. long, 2 mm. wide, ovate-rotund, 2-dentate, ciliate. Fructification not seen.
The armature of the leaf margins varies considerably in different plants, but the species can always be recognised by the sub-orbicular 2-dentate stipules, wide dorsal lobe and the axillary tufts of scale-like leaves.
On the ground in upland forests in both islands, also Stewart Island.
Schistochila repleta (Tayl.) Steph.
Jungermannia repleta Tayl., Hep. Nov. Zel. in Lond, Journ of Bot., 1844; Raoul, Fl. Novae Zeel. Gottschea repleta G., L. et N., Syn. Hep., 1844; Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., ii, 1867. G. macroamphigastria Col., G. heterocolpos Col., G. albistipula Col., G. pallescens Col., Trans. N.Z. Inst., Vol. xviii, 1885. G. gregaria Col., G. laciniosa Col., Trans. N.Z. Inst., Vol. xix, 1886. G. flavo-virens Col., Trans. N.Z. Inst., Vol. xx, 1887. G. guttata Col., G. longiseta Col., G. heterodonta Col., Trans. N.Z. Inst., Vol. xxi, 1888. Schistochila heterodonta Steph., Spec. Hep.
Plants medium, yellow-green. Stems 1–5 cm., usually about 2½, clothed most of the length with reddish-pink rhizoids, glabrous above. Leaves 4–5 mm., not soft in texture, yellow-green, brittle when dry, ovate-lanceolate, acute, sharply and coarsely toothed, basal ventral margin often jagged with large teeth, 3 or 4 short lamellae, dorsal lobe ⅓ shorter, semi-ovate, truncate, serrate, wing double, dorsal narrow with some teeth. Cells ca. 30 μ, basal 50–60 μ, with trigones. Sporophyte before breaking through the calyptra, enclosed in the involucre consisting apparently of the swollen hollow stem, overlaid at intervals with pairs of leaves and their stipules, and crowned with 2 bi-lobed lacerate leaves and a lacerate stipule. The calyptra which is 8 mm. long, is adnate with this involucre for its entire length.
Varies greatly in size and small forms have smaller toothing. A terrestrial species not uncommon on bush floors, also on rotting logs.
I have seen no plants from the South Island, but the Flora Novae Zelandiae records it from Port William, coll. Lyall.
Schistochila lehmanniana (Lindenb.) Steph.
Jungermannia lehmanniana Lindenb., Lehm. Pug. Gottschea hombroniana Mont., Annal de sc. nat., 1843; Tayl., Hep. Antarct., in Lond. Journ. of Bot., 1844. Gottschea lehmannia Nees, G., L. et N., Syn. Hep., 1844; Tayl., Fl. Ant., 1847; Mitt., Fl. Nov. Zel., ii, 1855; Fl. Tas., 1860; Handb. N.Z. Fl., ii, 1867. G. chlorophylla Col., Trans. N.Z. Inst., Vol. xviii, 1885. Schistochila lehmanniana Steph., Spec. Hep.; Rod., Tas. Bry., 1916.
Plants robust, pale green to pale yellow, in straggly clumps. Stems to 8 cm., branched, with rhizoids on the lower part, thick with paraphylls amongst the leaves and stipules. Leaves 5 mm. long, ovate to ovate-lanceolate, acute, soft in texture, crowded on the stems. One long lamella on the lower half, small or absent elsewhere, margins serrate with short sharp teeth, basal portion bare, dorsal lobe almost as long as the ventral, semi-ovate, acute, strongly arched from the base, armature very variable, usually serrate near the apex, and distantly ciliate at the base. Line of union reaching
about ⅔ the length of the leaf, wing double at the junction of the lobes, the ventral wider than the dorsal. Stipules 4-lobed, lobes convex with 2 or more divergent cilia at the apices, sinuses rounded, obtuse. Cells 30–40 μ, basal narrow, 60–70 μ long.
This species most resembles S. balfouriana in its softness and general appearance, but can be recognised by its sub-equal lobes, the dorsal not truncate, and the characteristic stipules. One very elegant form is from Jacksons Bay, in T. Kirk's collection, No. 4916 in the Plant Research Herbarium. The dorsal lobe is ciliated all round with cilia to 7 cells long, very acuminate with a long apical spine. Cells clear and somewhat larger than usual. This plant was identified by Stephani as S. marginata, but that identification is unquestionably incorrect. The direct opposite to this form is the following variety:
Var. compacta Hodg. nov.
Leaves generally shorter and broader, marginal teeth small and comparatively few. Stem paraphylls much less numerous.
This plant, No. 4920 Herb. Plant Research Bureau, is also in Kirk's collection, collected at Ulva, Stewart Island, undated.
Recorded in the Handbook from both islands, but I have seen no specimens from the North Island. It is possible that specimens of S. balfouriana have been mistaken for this plant. Also found in Tasmania.
Schistochila unguicularis (Tayl.) Steph.
Jungermannia unguicularis Tayl., Hep. Nov. Zel., in Lond. Journ. of Bot., 1844; Raoul; Fl. Nov. Zel. Gottschea unguicularis G., L. et N., Syn. Hep., 1844; Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., 1867. G. laete-virens Col.; G. simplex Col., Trans. N.Z. Inst., Vol. xviii, 1885. Schistochila unguicularis Steph., Spec. Hep.
Plants small to medium, pale green, overlaid in flat patches. Stems 1½–2½ cm., creeping, rhizoids along the greater part and scattered, small, bifid paraphylls on both dorsal and ventral surfaces. Leaves 3–4 mm. dull to pale green, contiguous, the lower spaced, oblong, obtuse, more or less decurrent, sub-entire, or serrations short and few, with a few basal cilia, lamellae present, sometimes faint, dorsal lobe small, ½ the length of the ventral, truncate, semi-ovate, more or less entire, line of junction generally straight, running towards the centre of the leaf; wing wide, single. Stipules 2-fid, dentate-ciliate, with or without lateral segments. Cells 30–40 μ with small trigones, basal 40–45 μ.
This species gradually merges into S. balfouriana. S. laete-virens (Col.) has a decurrent margin armed with a long tooth. Typical plants are rare. I have only one from the South Island, from Mount Grey, North Canterbury, coll. H. Hodgson.
Schistochila balfouriana (Tayl.) Steph.
Jungermannia balfouriana Tayl., Hep. Antarct., in Lond. Journ. of Bot., 1844; Raoul, Fl. Nov. Zeel.; Tayl., Fl. Antarct., 1847. Gottschea balfouriana, Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., ii, 1867; G., L., et N., Syn. Hep., 1844. G. nitida Col., G. trichotoma Col., G. marginata Col., Trans. N.Z. Inst., Vol. xviii, 1885. G. ciliistipula Col., Trans. N.Z. Inst., Vol. xix, 1886. G. truncatula Col., Trans. N.Z. Inst., Vol. xx, 1887. G. longiciliata Col., G. mitteniana Col., G. epiphyta Col., Trans. N.Z. Inst., Vol. xxi, 1888; G. marginata Steph., Journ. Linn. Soc., Vol. xxix, 1892; Schistochila balfouriana Steph., Spec. Hep.
Plants small to robust, pale to dull green, usually overlaid, forming mats. Stem 2–7 cm. simple or branched, with few or many paraphylls, red rhizoids numerous, sometimes in detached bunches at the bases of the stipules. Leaves 5–6 mm., soft, crowded towards the apex of the stem, often spaced on tall plants, with 3 or 4 toothed lamellae of varying lengths and heights, and traces of lamellae consisting of lumps at intervals, all running obliquely out to marginal incisions; obtuse, irregularly serrate, basal ventral margin armed with long and branched cilia, and usually with a large, lobate, irregularly ciliated serration; dorsal lobe semi-ovate, ½ to ⅓ as long as ventral, obliquely truncate, ventral margin serrate or ciliate-serrate, also the narrow dorsal wing below the junction of the lobes, the ventral wing usually serrate-ciliate, strongly so towards the base. Stipules sub-orbicular, deeply 4-fid, segments rounded-obtuse, margins recurved, fringed with long curved cilia. Cells 30–50 μ × 25 μ, basal 50–70 μ × 30 μ. Sporophyte sunk in the swollen stem, the overlaying leaves of which, including the uppermost, differ hardly at all from the ordinary foliage leaves, or narrowed to the apex and acuminate. Calyptra adnate with the walls of the involucre.
In this short paper and without numerous drawings, it is impossible to do justice to the very composite group represented by this and the preceding species. Moreover, they run into each other so that it is often impossible to decide between them, and yet the extremes are very different. In the Flora Novae Zealandiae, Mitten states concerning S. unguicularis, “… the smallest of the New Zealand species, allied to G. balfouriana, but differing in its much less dentate-ciliate leaves and stipules.” This is not of course entirely satisfactory. Perhaps it would be as well to consider the whole group as one variable species.
The two following examples will serve to show the variability in what is presumably S. balfouriana. One medium-sized plant from Omahanui, Wairoa, has apical serrations short, basal cilia long but few and simple, dorsal lobe very broad, sub-entire, abruptly narrowed to a truncate apex. While another very small plant from Waikaremoana has elaborately ciliated basal margins and serrations 3 cells long all round the much narrower dorsal lobe. In both of these the stipules are 4-fid and ciliated as in typical S. balfouriana.
The commonest of the species, found in all 3 islands in damp bush soil and coming down to low levels.
Schistochila muricata Hodgson et Allison sp. nov.
Planta sterilis, pallide flavo-virens, parva. Caulis 1–2 cm., simplex vel parce ramosus, nudus, radiculis rubris. Folia ca. 45 mm. × 15 mm., imbricata, ovato-lanceolata, muricata, acuta, marginibus serratibus; lobus anticus brevior quam posticus, obliquiter truncatus, similiter armatus et muricatus. Amphigastria bifida ad medium, lobis cum uno laterali, brevi segmento, omnibus marginibus spinuloso-dentatibus. Areolae obscurae, basales, ca. 40–60 μ X, ca. 20–30 μ.
This plant has a creeping habit similar to that of S. unguicularis, and is much the same size; but even without a lens, the short, hard, sloping points which cover the foliage give it a distinctive look. On the upper parts of the stems the leaves are imbricate and horizontal,
while toward the base they become scarcely contiguous and less horizontal. Red rhizoids are present on the lower half of the stem on the ventral side, but there appear to be no paraphylls. The characteristic narrow-lanceolate leaves are without lamellae, though traces are sometimes discernible; narrow towards the apex, acute, scarcely acuminate, muricate, the margins closely and irregularly serrate, the teeth hyaline, resembling the processes on the lamina, straight, curved, or sub-hamate. The dorsal lobe, which is narrowly semi-ovate and obliquely truncate, extends to a little more than half the length of the ventral, the toothed line of insertion running obliquely towards the middle of the leaf. The stipules, which have smooth surfaces, are imbricate on the upper part of the stem, sub-imbricate to contiguous on the lower, about 1 mm. in length or a little more, and more than half the length in width. The lobes are sometimes with a short lateral segment on each outer margin, all segments being acuminate with margins that are spinulose-toothed, and not always flat. The cells of the stipules are thick-walled, and about 50 μ, long in the undivided part, and gradually become much smaller, from 20–30 μ, in the segments. The cells of the leaves are obscure and opaque owing to the processes, but the basal are hyaline, about 40–60 μ long by 20–30 μ wide. In both gatherings the plants are sterile.
This interesting plant was discovered by Mr. V. D. Zotov by a watercourse north of Field Hut, Tararua Mountains, ca. 2600 ft., January 1, 1934. Early in 1935 it was gathered by Messrs. G. Simpson and J. S. Thomson in the watershed of Kaituna Creek, Nelson.
Both of these co-types are in the Herbarium of the Plant Research Bureau, Wellington.
Schistochila glaucescens (Hook.) Steph.
Jungermannia glaucescens Hook., Musci Exotici, 1818. Gottschea glaucescens Nees; G., L. et N., Syn. Hep., 1844; Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., ii, 1867. Schistochila glaucescens Steph., Spec. Hep.
Plants medium or robust, of straggly growth in loose cushions, pale yellow to blue-green. Stems 3–8 cm., much branched, clothed with ciliate paraphylls, red rhizoids at base. Leaves 2½–5 mm. according to size of plant, pubescent, oblong, pinnatifid, with numerous erect lamellae margined with branched hairs, scattered branched hairs on inside or dorsal surface, dorsal lobe strongly arched, ½ the length of ventral, truncate with numerous parallel lamellae fringed with jointed hairs on the dorsal side; smooth on the inside or ventral surface. Stipules orbicular, 4-fid, with long, branched cilia in rows and fringing the margins. Cells 6-sided, ca. 30–40 μ, basal 60–70 μ × 20–30. Cells of stipules ca. 60–80 μ × 25. Involucre oblong-cylindrical, clothed with leaves and stipules, and surmounted with a pair of leaves with their stipule, all joined at their bases.
Easily recognised by the pale colour and the pubescent foliage and stipules and the only New Zealand plant with a lamellate dorsal lobe.
On ground or rotting logs under bush or scrub, but not at low levels except on Stewart Island. There are 4 gatherings of this plant in Mr. V. D. Zotov's fine collection from the Tararuas.
Schistochila ciliata (Mitt.) Steph.
Gottschea viliata Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., ii, 1867. G. chlorophylla Col., Trans. N.Z. Inst., Vol. xviii, 1855. Schistochila viliata Steph., Spec. Hep.; Rod., Tasman. Bry., ii, 1916.
Plants usually in dense low cushions, pale green, lighter at the tips, often tinted purplish or rose. Stems 2–7 cm. simple or branched, thick, rhizoids at the base, paraphylla present on robust plants. Leaves 5 mm. long, 3·5 mm. broad, densely imbricate, without lamellae, broadly ovate, margin undulate, fringed all round with long 5-celled hair-like cilia; dorsal lobe almost as long, ciliate, a double wing below the line of union; ventral wing ·25 mm. wide, cilia shorter and fewer towards the base; dorsal much narrower, strongly ciliate. Stipules large, rounded, 4-lobed, the 2 middle ones larger, all armed with cilia. Cells 30–40 μ with trigones, basal scarcely larger. Sporophyte very deeply immersed in cylindrical involucre which is clothed with leaves and stipules, surmounted with 2 very wide leaves and a stipule, with sterile archegonia at the edge of the calyptra mouth.
Neither Mitten nor Stephani mentions paraphyllia on the stem, but they are quite numerous on robust plants from Stewart Island, coll. Mrs. J. D. Smith.
On damp forest floors, rotting logs and under scrub on higher levels throughout New Zealand, also Tasmania.
Schistochila splachnophylla (Tayl.) Steph.
Jungermannia splachnophylla Tayl., Hep. Antarct., in Lond. Journ. of Bot., 1844. Gottschea splachnophylla G., L. et N., Syn. Hep., 1844; Tayl., Fl. Antarct., 1847; Mitt., Fl. Nov. Zel., ii, 1855; Handb. N.Z. Fl., 1867. Schitochila splachnophylla Steph., Spec. Hep.
Plants medium, when dry, brown in colour and deeply set in the substratum like Riccia. Stems creeping, excessively thick and fleshy, thick with rhizoids impregnated with detritus. Leaves 3 mm. obliquely spreading, entire, thick, fleshy, truncate-obtuse imbricate; the dorsal lobe semi-cordate, truncate, obtuse, sub-equal to the ventral and placed on it at about an angle of 45°, with a broad ventral wing as a keel, which neatly fits into the space between the lobes of the leaf below it. Stipules ovate-oblong, shortly 2-fid, sinus small and rounded, with numerous rhizoids arising from the fleshy cushion-like centre. Fructification unknown.
In the Flora Novae Zealandiae Mitten says of this plant: “…In the younger stems the leaves vary considerably, and are found with the ventral lobe ovate-acute, the dorsal also ovate with one or two obtuse teeth. The free portion below the line of union of the 2 lobes is also wider. The stipules in the larger plants are somewhat of a semi-orbicular form, their apices obtuse; in the smaller plants they are oblong and bifid.”
From the description of Gottschea pachyphylla in the Synopsis Hepaticarum it would seem that this also is S. splachnophylla.
S. splachnophylla was first discovered at Cape Horn and the Straits of Magelian by Mr. Menzies, while the New Zealand plants were first found on the Ruahines by Colenso. I have heard of no
other gathering till Mr. V. D. Zotov found it in a watercourse on Mount Hector in the Tararuas. Seeing it is a South American plant, one would expect it to turn up on the mountains of the South Island.
Schistochila kirkiana Steph.
Plants small, diœcious, yellow-green. Stems creeping, 1–2 cm., overlaid in mats, rhizoids red or yellow-green, no paraphylls seen. Leaves 2·8 mm. long, yellow-green, obliquely spreading, ovate-lanceolate, crowded on the upper part of the stem, strongly spinose-serrate along the ventral margin and part of the dorsal margin, with an apical spine which is 6 cells long; dorsal lobe sub-equal in length, wider at the arch, semi-ovate, margin spinose-serrate with a triangular lobe at the base, upper part free; line of union distant ⅓–½ the width of the leaf from the dorsal margin, thus forming a very wide wing, with a narrow scarcely toothed wing of its own. Stipules small, bifid. Cells ca. 40 μ with nodulose trigones, basal ca. 50–70 μ × 35μ. Antheridia 2–5 in axils of upper leaves. Sporophyte black 1·8 mm. long embedded in the tubular involucre, overlaid with leaves with spiny apices, the unruptured apex of the involucre, i.e. the free part of the calyptra, rough with prominences.
What appears to be an aquatic form of this species was collected in Regina Valley, South Westland, by Mr. B. Teague. It is a much laxer and taller plant, leaves twice the ordinary size and more widely spaced along the stem, grey-green, soft, often with a drooping ventral lobe, the narrow secondary wing strongly toothed, and the whole plant a mass of sandy detritus.*
Still another form is from the National Park, collected by Mr. G. O. K. Sainsbury. This is also grey-green, with leaves imbricate and less patent, giving the stem a subterete appearance. The rhizoids are excessively thick on the stem, imparting to the whole plant a pinky tinge when wet. I noticed 2 rhizoids attached to the ventral surface of a leaf. This gathering was also from a sandy habitat and very hard to clean.
Of the 4 other plants in my collection 3 were collected in the Rotorua and Murupara districts by Mr. K. W. Allison and one from Arthurs Pass was collected by Miss P. Milligan. In this plant the rhizoids are hyaline-yellow-green, and the dorsal lobes of the leaves directed upwards, which gives the plants a prickly appearance, more so than in Mr. Allison's plants.
Schistochila colensoana Steph.
Plants very small, creeping, yellow-green. Stems ½–¾ cm., thickly clothed on the ventral side with long hyaline rhizoids. Leaves
[Footnote] * Since writing the above, I have received from Miss L. B. Moore a plant collected at Wilkin River, Lake Wanaka, October, 1941, by Mr. G. Simpson, which is a luxuriant form of the Regina Valley plant. It has the same broad, soft, though yellow-green leaves, with a drooping ventral lobe; but the margins of the lobes are softly ciliate and not spinous-toothed. The stems are 5 cm. tall and in some cases branched, and the whole plant is very far removed in appearance from the little prostrate form. Nevertheless, the fundamental structure and the nodulose areolation are the same in all three plants, and I cannot bring myself to separate them.
crowded on the stem, 2½ mm. long, ovate-elliptical, obliquely truncate, with scattered teeth on the ventral margin and around the apex; dorsal lobe sub-equal, joined to the ventral for almost its entire length, broadly semi-ovate, obliquely truncate with an arista at the upper corner, entire or sub-entire; wing broad as in S. kirkiana. Stipules small and difficult to pick up amongst the rhizoids, orbicular-ovate, bifid to nearly halfway, segments obtuse. Cells about 40 μ. Involucre surmounted by a pair of toothed leaves with their stipule. Sterile archegonia on the ruptured apex of the calyptra. Capsule valves 2½–3 mm. long, ¼ as broad.
The above is a description of a plant collected by Mr. Allison, between Taupo and Rangitaiki, ca. 2500 ft. It was seen by Prof. Herzog, who pronounced it to be a minor form of Stephani's S. colensoana, and we agree with this identification, though Stephani's description says that stipules are absent from his plant, and this definitely has small ones. They may have been overlooked by Stephani. As a matter of fact, this seems to be either very closely related to, or a form of S. kirkiana. However, the armature of the leaves is different, and perhaps the line of insertion of the dorsal lobe, so at any rate until fresh plants are found, it must remain a separate species.
Schistochila pinnatifolia (Hook.) Steph.
Jungermannia pinnatifolia Hook., Musci Exotici, 1818. J. ciliigera Tayl., Hep. Antarct., in Lond Journ. of Bot., 1844. Gottschea pinnatifolia Nees, G., L. et N., Syn. Hep., 1844; Tayl., Fl. Antarct., 1847; Mitt., Fl. Nov. Zel., 1855; Handb. N.Z. Fl., 1867. G. plumosa Col., Trans. N.Z. Inst., Vol. xx, 1887. Schistochila pinnatifolia St., Spec. Hep.; Rod., Tas. Bry., 1916.
Plants robust or medium, in tufted cushions, green to golden-brown. Stems 3–7 cm., sparingly dichotomously branched, flexuous, apices sometimes nodding or circinate. Leaves 2–3 mm. long, con-duplicate, bi-lobed to ¾ the length of leaf, imbricate, lobes sub-equal, incised-pinnatifid, the dorsal inflated; all margins fringed with glistening cilia, also the narrow wing below the junction of the lobes. Stipules 0. Areolation distinct, with small rounded cells and large trigones. Cells 20–30 μ, basal oval ca. 50 μ. Fructification not seen, but Hooker in the Musci Exotici states that the upper half of the calyptra is free from the involucre, bearing sterile archegonia at its apex, while the involucre is 4-fid, the segments being incised-pinnate and ciliate.
A very beautiful plant found throughout New Zealand in upland forest, also in the Auckland Islands and Tasmania.
Schistochila tuloides (Tayl.) Steph.
Jungermannia tuloides Tayl., Hep. Nov. Zel., in Lond. Journ. of Bot., 1844. Gottschea tuloides G., L. et N., Syn. Hep., 1844; Mitt., Fl. Nov. Zeol., 1855; Handb. N.Z. Fl., 1867. Schistochila tuloides Steph., Spec. Hep.
Plants medium, sometimes corticolous, pale green to cream. Stems 2 cm. procumbent, simple or branched, apices sometimes branched, clusters of archegonia at intervals along the upper part of the stem, surrounded by small, irregularly shaped, sometimes bifid leaves, often with a long narrow basal segment. Leaves 4 mm., imbricate, oblong-lanceolate, sub-entire to irregularly serrate, the
armature varying greatly, even on the same plant, papillose, specially in the upper parts; dorsal lobe sub-equal in length, variable in size and width, margins reflexed, line of insertion extending from ⅔ to ¾ the length of the leaf, upper part free, no secondary dorsal wing. Stipules 0. Cells of unequal length, ca. 30–40 μ, trigones large, basal cells to 50 μ with somewhat sinuous walls, trigones very large; papillae not always present, sometimes reduced to a convexity. Involucre overlaid with leaves that are adnate in the lower part, uppermost leaves irregular in shape, sometimes lacerate, mouth of calyptra with sterile archegonia.
This is the only species in which I have observed groups of archegonia on the stem, which must be unusual in a genus with a terminal fructification. Also unusual is the (perhaps occasional) habitat, which is on bark. Miss L. B. Moore states that on Mount Moehau it is “fairly common, usually on the underside of sloping kauri branches.” Mr. V. W. Lindauer also sends it from the Waipoua Kauri Forest. Also at the base of forest trees. I have seen only two gatherings from the South Island, one collected by Messrs. Simpson and Thomson, at Lead Hills, Bainham, Nelson, and the other from Taylorville, Westland, coll. M. Berry.
This is another of Colenso's notable discoveries, easily recognised by its hard looking pale foliage, the sub-equal lobes and the absence of stipules.
Schistophylla conchophylla Hodgson et Allison sp. nov.
Planta sterilis, singularis, brunnescens, erecta vel flexuosa. Caulis ad 6 cm. simplex vel raro ramosus, ubique nudus. Folia ca. 3 mm., imbricata, late ovata, bi-loba, inflata, elamellata; lobus anticus fere aequalis, semi-cordatus, margine integro vel paucidenticulato. Alae ·3 mm. latae. Amphigastria 0. Areolae ca. 20–30μ, basales ca. 40–50 μ × 25 μ, trigonis ubique maximis. Fructus ignotus.
The name of S. conchophylla was suggested for this plant by Prof. Herzog, and it is very apt, as the sub-equal, strongly inflated leaf-lobes bear a strong resemblance to a bi-valved shell; especially when wet. The plants are brownish-green to varying shades of brown, all separate in the packet, mixed with stems of Rhacocarpus australis. The stems are more or less flexuous, and bare of either paraphylls or rhizoids. The leaves are imbricate, obliquely set on the stem, and broadly ovate, with 3 or 4 small teeth on the antical and postical basal margins, and scattered teeth along the margin of the dorsal lobe. The line of union of the lobes reaches at most about ¾ the length of the leaf, leaving the remaining quarter free. The concave lobes are more nearly equal than in any other species and possess a characteristic sheen. The cells also are typical, being rounded, with thin walls, and with large confluent trigones, becoming oval towards the base and still with trigones. The areolation recalls that of S. pinnatifolia, but in that plant the cells are much more uneven in size and have thicker walls. S. conchophylla resembles S. tuloides in its concave lobes, sub-entire margins, and absence of stipules, but that plant is very pale in colour and has a creeping habit.
So far this new species has been found in only one locality, viz., Mount Stoke, Marlborough, where it was collected by Mr. J. H. McMahon, the date unspecified.
The type is now in the Herbarium of the Plant Research Bureau, Wellington.
It is interesting to note that of the 15 species described above, 4 were discovered by Mr. Menzies at Dusky Bay in 1791. They were S. pinnatifolia, S. nobilis, S. glaucescens, and S. appendiculata, all described and figured in Sir W. Hooker's Musci Exotici, in 1818.
Notes on Colenso's Species.
In earlier numbers of the Trans. N.Z. Inst., Rev. W. Colenso has described the following species of Schistochila:
|S. compacta (Col.)||Trans., 1883, Vol. xvi, p. 349|
|S. laetevirens (Col.)||1885, Vol. xviii, p. 238|
|S. nitida (Col.)||1885, Vol. xviii, p. 238|
|S. macroamphigastria (Col.)||1885, Vol. xviii, p. 238|
|S. heterocolpos (Col.)||1885, Vol. xviii, p. 239|
|S. trichotoma (Col.)||1885, Vol. xviii, p. 240|
|S. chlorophylla (Col.)||1885, Vol. xviii, p. 240|
|S. bicolor (Col.)||1885, Vol. xviii, p. 240|
|S. pallescens (Col.)||1885, Vol. xviii, p. 241|
|S. marginata (Col.)||1885, Vol. xviii, p. 241|
|S. albistipula (Col.)||1885, Vol. xviii, p. 242|
|S. simplex (Col.)||1885, Vol. xviii, p. 242|
|S. ramulosa (Col.)||1885, Vol. xviii, p. 243|
|S. ciliistipula (Col.)||1886, Vol. xix, p. 284|
|S. laciniosa (Col.)||1886, Vol. xix, p. 284|
|S. truncatula (Col.)||1887, Vol. xx, p. 247|
|S. flavo-virens (Col.)||1887, Vol. xx, p. 248|
|S. squarrosa (Col.)||1887, Vol. xx, p. 248|
|S. plumulosa (Col.)||1887, Vol. xx, p. 249|
|S. homophylla (Col.)||1887, Vol. xx, p. 250|
|S. guttata (Col.)||1888, Vol. xxi, p. 53|
|S. longiciliata (Col.)||1888, Vol. xxi, p. 54|
|S. longiseta (Col.)||1888, Vol. xxi, p. 54|
|S. heterodonta (Col.)||1888, Vol. xxi, p. 55|
|S. stenocarpa (Col.)||1888, Vol. xxi, p. 55|
|S. mitteniana (Col.)||1888, Vol. xxi, p. 56|
|S. moniliformis (Col.)||1888, Vol. xxi, p. 56|
|S. epiphyta (Col.)||1888, Vol. xxi, p. 57|
|S. winklemannii (Col.)||1888, Vol. xxi, p. 57|
|S. clandestina (Col.)||1889, Vol. xxii, p. 454|
Of these I have seen the following: S. nitida, S. laetevirens, S. simplex, S. guttata, S. longiseta, S. pallescens, S. marginata.
These descriptions in the Transactions make interesting reading. Minor differences seemed very important to Colenso, and indeed they might well be, but at the same time he could not appreciate the fundamental resemblances which band a group of plants together under one specific name. Moreover, he could not recognise his species a second time and thus establish them, for the majority of plants in the Colenso collection have no names at all.
Fortunately, Dr. Fr. Stephani overhauled Colenso's material at Kew and published his results in the Journal of the Linnean Society, Vol. xxix, 1892. He allowed 5 of these species to stand, remarking
Figs. 1–3—S. unguicularis; Mount Grey, Coll. H. M. Hodgson; Fig. 1, Leaf; Figs. 2–3, Stipules. Figs. 4–5—S. nitida, ex Herb. Col. = S. balfouriana; Fig. 4, Leaf; Fig. 5, Stipule. Figs. 6–7—S. repleta, ex Herb. Col.; Fig. 6, Leaf; Fig. 7, Stipule. Figs. 8–9—S. tuloides; Fig. S, Leaf; Nelson, Coll. Messrs. Simpson and Thomson; Fig. 9, Leaf; Waipoua Kauri Forest, Coll. V. W. Lindauer. Figs. 10–11—S. lehmanniana; Stewart Island, Coll. Mrs. J. D. Smith; Fig. 10, Leaf; Fig. 11, Stipule with Paraphyll. Fig. 12—S. lehmanniana var. compacta; Leaf and Stipule; Ulva, Stewart Island, ex Herb. T. Kirk. Figs. 13–14—S. splachnophylla; Tararua Mountains, Coll. V. D. Zotov; Fig. 13, Leaf; Fig. 14, Stipule. Figs. 15–16—S. conchophylla; Mount Stoke, Nelson, Coll. J. H. McMahon; Fig. 15, Leaf; Fig. 16, cells, ca. 200 X. Figs. 17–18—S. appendiculata; Kapiti Is., Coll. Students, per Mr. McKenzie; Fig. 17, Leaf. Fig. 18, Stipule.
Fig. 19—S. pinnatifolia, Leaf, with drooping ventral lobe. a; L. Waikare-iti, Coll. E. A. Hodgson. Figs. 20–22—S. kirkiana; Arthurs Pass, Coll. P. Milligan; Fig. 20, Stem with Leaves and Stipule, ventral; Fig. 21, Leaf, dorsal; Fig. 22, Stipule. Figs. 23–24—S. kirkiana, aquatic form; Regina Valley, South Westland, Coll. B. Teague; Fig. 23, Leaf with drooping ventral lobe, b; Fig. 24, Stipule. Figs. 25–27—S. colensoana; between Taupo and Rangitaiki. Coll. K. W. Allison. Figs. 25–26, Leaves; Fig. 27, Stipule. Figs. 28–30—S. nobilis; Haast Pass, Coll. Miss Matthews; Fig. 28, Leaf; Fig. 29, Stipule; Fig. 30. Tuft of Paraphylls reduced to 2. Fig. 31—S. ciliata Leaf; Waikaremoana, Coll. E. A. Hodgson. Figs. 32–33—S. muricata; Tararua Mountains, Coll. V. D. Zotov; Fig. 32, Leaf; Fig. 33, Stipule.
that they “appear valid.” These 5 were S. marginata, S. heterodonta, S. guttata, S. ramulosa and S. squarrosa. I have copies of his drawings of S. marginata, S. heterodonta, and S. ramulosa, in the Icones Hepaticarum, and without any doubt
S. marginata (Col.) is S. balfouriana (Tayl.)
S. heterodonta (Col.) is S. repleta (Tayl.)
S. ramulosa (Col.) is S. tuloides (Mitt.)
S. guttata (Col.) is also S. repleta (Tayl.), and from the description, S. squarrosa (Col.) must be S. nobilis (Hook.), as this plant is too outstanding for any minor variation to constitute a separate species.
It is extraordinary that Stephani should have overlooked these older established species, when he allowed Colenso's names to stand. Other species of Colenso's he reduced as follows:—
[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]
|S. laetevirens (Col.)|
|S. nitida (Col.)|
|S. truncatula (Col.)|
|S. trichotoma (Col.)|
|S. simplex (Col.)||is S. marginata (Col.)|
|S. longiciliata (Col.)|
|S. ciliistipula (Col.)|
|S. epiphyta (Col.)|
|S. dichotoma (Col.)||is S. squarrosa (Col.)|
|S. winklemanii (Col.)||is S. appendiculata (Hook.)|
|S. moniliformis (Col.)|
|S. longiseta (Col.)|
|S. clandestina (Col.)|
|S. heterocolpos (Col.)|
|S. gregaria (Col.)|
|S. laciniosa (Col.)||is S. heterodonta (Col.)|
|S. macroamphigastria (Col.)|
|S. flavo-virens (Col.)|
|S. pallescens (Col.)|
|S. plumulosa (Col.)||is S. pinnatifolia (Hook.)|
|S. chlorophylla (Col.)||is S. ciliata (Mitt.)|
Of these, S. laetevirens and S. simplex are S. unguicularis. The fact that Stephani included them in the group which he reduced to S. marginata, i.e. balfouriana, suggests, as I mentioned before, that it might be feasible to consider these two plant groups as one.
It might be as well to mention that, with regard to S. marginata, Colenso must have had a mixture here, as the duplicate in the Napier collection is S. repleta, a different plant from Stephani's drawing in the Icones Hepaticarum.
It is evident from the above lists that the following plants were not seen by Stephani: S. compacta, S. bicolor, S. albistipula, S. homophylla, S. steno-carpa, S. mitteniana. A study of the descriptions, combined with a working knowledge of the established species, leads one to the following conclusions:—
S. compacta (Col.) seems to be S. colensoana Steph.
S. bicolor (Col.) seems to be S. repleta (Tayl.)
S. albistipula (Col.) seems to be S. repleta (Tayl.)
S. homophylla (Col.) seems to be S. tuloides (Mitt.)
S. mitteniana (Col.) seems to be S. balfouriana (Tayl.)
S. steno-carpa (Col.) seems to be S. repleta (Tayl.)
Of these, S. compacta is the most interesting. The description says: “Leaves semi-circular, broadly elliptic and sub-quadrate in outline, margin entire, decurrent, sometimes very sparingly toothed towards the base.” This could apply to the leaves of S. colensoana, and, if so, there would be another locality, viz., near Norsewood, for that rare plant.
In conclusion, I wish to thank the authorities of the Dominion and Hawke's Bay Museums for the loan of Colenso's collection, Dr. H. H. Allan and Mr. V. D. Zotov for the use of the collections in the Plant Research Herbarium, and the collectors who so kindly send me plants.
Hooker, Sir William, 1818. Musci Exotici. London.
Gottsche, C. M., Lindinberg, J. B. G., et Nees, C. G., ab Esenbeck, 1844. Synopsis Hepaticarum. Hamburg.
Hooker, Sir J. D. Botany of the Antarctic Voyage—
Dr. Taylor, in Flora Antarctica, London, 1847.
W. Mitten, in Flora Novae Zealandiae, Part ii, London, 1855.
W. Mitten, in Flora Tasmaniae, London, 1860.
Mitten, W. In Sir Joseph Hooker's Handbook of New Zealand Flora, Part ii, London, 1867.
Colenso, Rev. W. Descriptions of newly-discovered Cryptogamic Plants, Trans. N.Z. Inst., vol. xvi, p. 349, 1883; vol. xviii, pp. 238–243, 1885; vol. xix, p. 284, 1886; vol. xx, pp. 247–250, 1887; vol. xxi, pp. 53–57, 1888; vol. xxii, p. 454, 1889.
Stephani, Fr., Species Hepaticarum, in part; nos. and dates of vols. not stated.
—— Icones Hepaticarum, in part; nos. and dates of vols. not stated.
—— Colenso's New Zealand Hepaticae, Journ. of Linn. Soc., vol. xxix, pp. 263–280, London, 1892.
Rodway, L., 1916. Hepatics, Tas. Bry., vol. ii, Hobart.
Buch, H., 1939. Die Schistochila-Arten der Inseln Sumatra, Java and Celebes, Ann. Bry., vol. xii, pp. 1–20, Leiden.