New Zealand Hepaticae (Liverworts)—XIII

Transactions of the Royal Society of New Zealand : Botany, Volume 1, Issue 12, 16 May 1962, Page 139

New Zealand Hepaticae (Liverworts) —XIII

E. Amy Hodgson

K. W. Allison

A. Review of the New Zealand Species of the Genus Temnoma and of Anoplostoma, a new Genus

By

and

[Received by the Editor, April 12, 1961.]

Summary

Description of genus Temnoma, key to species of Temnoma, description of seven species including one new species, T. schusteri Hodgson & Allison, and one new combination. T. palmatilobum (Hodgson) comb, nov., other species being T. pulchellum, T. quadrifidum, T. quadripartitum, T. palmatum and T. corrugatum.

Description of new genus Anoplostoma with 2 species described as 2 hew combinations, Anoplostoma vaginatum (Herzog) comb, nov., type, and Anoplostoma whiteleggei (Carr. & Pears.) comb, nov., in 2 new subgenera, subgenus Anoplostoma and subgenus Trigona.

New Zealand, which may be considered the centre of distribution of the genus Temnoma, has 7 species of a probable total of 8. A detailed study of these New Zealand series reveals that vegetatively Temnoma has a wide range of sometimes conflicting characters, and the concept of the genus (Schuster, 1959) has accordingly been extended to accommodate these characteristics.

We extend grateful thanks to the late Dr Marshall Howe for the loan of the type of Temnoma quadriftda Mitt., to Dr Persson for a piece of the type of T. dusenii (S.teph.) Schuster, and a specimen of T. quadripartitum (Hook, senr.) Mitt., from Patagonia, and for literature; to Mrs J. Winkley, secretary of the Hawke’s Bay Branch of the Royal Society of New Zealand for forwarding literature from the library, also to Dr Th. Herzog for the name Anoplostoma.

Temnoma Mitt.

Temnoma Mitt., Handb. N.Z. FL, ii, 753, 1867, Supp.

Plants dioicous, or monoicous, small to robust, creeping or erect, in pure colonies or with other hepatics, from low to alpine levels. Stems simple or sparely branched, branches normally of the Frullania type; with the branch replacing the ventral half of, a lateral leaf, intercalary branching occasionally present, monopodia!, cortical cell-walls incrassate in T. corru-

gaturrr, rhizoids rare or plentiful, in tufts at the base of an underleaf or branch, truncate or with apices swollen into a shapeless mass of cells which may again produce rhizoids. Gauline leaves 4-fid, or 4-par,tite, mainly 3-partite in T. schusteri, bis-bilobed in T. palmatiloba, margins entire or spinous-ciliate, insertion succubous to transverse, imbricate to distant; underleaves usually somewhat smaller than the lateral, and similarly lobed and armed, sometimes bilobed in T. corrugatum. Cell-

type varied, usually quadrangular, square to oblong with thinnish walls, but thick-walled in T. corrugatum and collenchymatous in T. schusteri. Floral leaves large, 4-lobed, with toothed or spinous margins. Perianth sub-cylindrical, 3-gonous when fresh, mouth not contracted when the perianth is empty, sub-floral innovations present in T. pulchellum. Capsule firm, dark red-brown, walls and seta thick, spores small. Androecia usually terminal, bracts conduplicate, folded from the middle sinus, spinescent. No ventral $ bracts have as yet been found in this genus.

Gemmae not known.

Investigations by Dr R. M. Schuster show that the type of Temnoma, T. pulchellum (Hook, senr.) Mitt, has a thick capsule wall (4-5 stratose) and a thick seta 9-10 cells across. Whether these characteristics obtain for T. quadripartitum (Hook, senr.) Mitt., T. corrugatum (Steph.) Schuster, T. palmatilobum (Hodgson) Hodgson & Allison, T. schusteri Hodgson & Allison is not known.

Rhizoids with swollen, misshapen tips have been observed in T. palmatum, T. palmatilobum, T. corrugatum, and T. schusteri.

Temnoma differs from Blepharostoma to which the species have at one time or another been consigned, in the broader leaf-lobes from a basal lamina of varying depth. In Blepharostoma the leaves are divided almost to the base into 2-5 hair-like segments, composed of a single series of cells throughout. The wide, truncate perianth is also characteristic of Temnoma but not of Blepharostoma, while the latter has a 2-stratose capsule wall.

Although the genus has been re-defined to accommodate species and characters unknown to Schuster (1959) the family Blepharostomaceae still seems adequate for its inclusion.

Key to Species

1. Plants small, leaves and underleaves equal in size, tripartite, cells with thick, bulging walls (collenchymatous) T. schusteri Plants small to robust, leaves 4-lobed, underleaves mostly somewhat smaller, cells quadrangular with thin walls (except in T. corrugatum) 1 2

2. Leaves twice lobed with the middle sinus deeper, leaf-lobes obtuse and margins quite entire except in the floral leaves, insertion transverse ...... ...... ...... T. palmatilohum Leaves not twice lobed, sinuses mostly of even depth, cauline leaf-lobes with margins and apices either toothed or spinous (rarely so in T. quadrifida) ...... 3

3. Marginal spines curved, affording a unique appearance, cells of leaves and stem cortex incrassate with oval cavities, underleaves occasionally only bifid T. corrugatum Marginal and apical spines when present, straight, leaf and cortical cells quadrangular with thin to thinnish walls 4

4. Plants small, leaves deeply 4-partite with basal lateral teeth oppositely placed, transverse, lobes narrow-elongate, widely diverging, cells small, not elongated, monoicous in 1 South American plant „.... ...... ...... ...... ...... ...... T. quadripartitum Leaves 4-lobed to i or less, succubous ...... 5

5. Leaves 4-lobed to about halfway, lobes lanceolate, apices produced to a point or a short spine, usually 2-3. cells long T. quadrifidum Leaves with spines from 5-7 cells long in a single series, spines golden, glossy, leaves succubous 6

6. Plants medium to robust, leaves 4-lobed with both marginal and apical spines, perianth long and slender T. pulchellum Plants small, leaves with short lobes and shallow sinuses, lobe apices tipped with spines with occasional lateral, smaller spines on the margin of the leaf-base or lamina, perianths wide and shortish with a truncate mouth .; T. palmatum

Temnoma pulchellum (Hooker senr.) Mitt. (Text-fig. 1, figs. 21-22) Jungermannia pulchella Hook, senr., Musci Exotici Tab. 114, 1818. Temnoma pulchella Hook, senr., Mitt, in Hook. Handb. N.Z. FI., 11, Supp., 750, 1867. Blepharostoma pulchellum (Hook, senr.) Steph. Spec. Hep., 3, 640, 1909.

Plants dioicous, terrestrial or lignicolous, in pure associations or with other bryophytes, beautiful with glossy spines, golden brown to dull green. Stems to as much as 3.5 cm, but rarely more than 2 cm long, 0.3 mm broad, more or less prostrate, a little flexuous, sparsely branched. Leaves variable as to lobing, succubous in varying degrees, closely set on most stems, obliquely to horizontally spreading, broader than tall, expanding from a narrow base with more or less straight sides, ca. 1.1 mm broad and 0,5 mm tall excluding the spines, usually 4-lobed, with perhaps 2 lateral sub-basal lobules; sinuses variable in depth, ca. 0.2 mm, obtuse, about as broad as the lobes, margins reflexed; lobes with 4-6 irregularly placed mostly apical, divaricating spines with one or two lateral spines; spines or setae articulated, a little swollen at the joints, consisting of 5-7 elongated cells, ca. 50//, long and 25 jx wide, the setae gradually tapering upwards. In the sinuses of the lobes, any spines present may be displaced from the margin; cells ca. 23//,, hexagonal or subhexagonal in the upper portion, midleaf lengthening to ca. 45//, long and 25//, broad, basal, narrow-elongate to 70//, long, the midleaf and basal cells arranged in tiers, walls moderately thick, trigones absent or minute, cuticle smooth. Underleaves 4-6-lobed, reniform with arched insertion, similar to the lateral leaves as regards spines and cells. Floral leaves in 3-4 rows, increasing in size upwards, the 2 upper pairs not closely imbricated and with the lower portion appressed to the perianth, deeply lobed and ciliated. Perianth to nearly 4 mm, straight or decurved, flimsy, often sterile, narrowed to the mouth, trigonous, mouth deeply lobed, lobes bi-tri-fid margins and apex longly setose, cells hyaline, very elongated to 100/x or more, narrow. Spores golden, small, ca. 8//,, very numerous, elaters short, 0.1-0.2 mm, bispiralled. Capsule obovate, dark reddish brown, 1.3 mm long x 0.7 mm wide. $ bracts to 8 pairs, at the upper portions of the stem and branches, saccate, the inflated portion reaching the length of the ventral margin, the lamina 3-lobed, each lobe with 3 setae, the middle one the longest; autheridia single, 0.2 mm in diameter.

There are intergrading forms between this species and T. palmatum, with the lobes decreasing in size till on some weak stems almost non-existent, the leaf consisting of the lamina and the spines. One specimen from Rakiahua forest, Stewart Island 9813 W.M. has the spines very strongly developed with some even branched at the base.

This species, which is most distinct and attractive looking, is easily recognized by the lobed leaves, each lobe crowned with straight, articulated, glossy spines. It is similar in appearance to Trichocoiea lanata, but the golden colour distinguishes it.

Rodway (1916) describes Tasmanian plants as having the cuticle of the leaf upper cells asperate, and the lower ones striate.

Though not common, the species is distributed throughout New Zealand.

North Island; Great Barrier Island, with the rare Radula dentata T. Kirk; Bay of Islands area, with Trichocolea lanata 7753,; between Dargaville and Opononi 1,200 ft, 652; Horokino Bush, Mangapehi, 5561, V. W. Lindauer; Waipoua Forest, on rotting logs, H 707, H 660, H 657, K.W.A.; bog, Aniwaniwa V. Urewera National Park, 9314, H. M. Druce; damp bushed gully, Levin, 630, L.8.M.; Herb. Mitten coll. Kerr; base of Coromandel Pen., H 5218, J. M. Mitchell; Coromandel Pen., H 1442, L. J. Matthews; also Nos. HISBO, H 1583, H 1582, HISBI, Waipoua Forest, K.W.A.

South Island: Moderate shade amongst grass, forest remnant, Taieri Mouth, 902; bush track, Swampy Hill, Dunedin, 9629; roadside banks, Lake lolanthe, S. Westland, 5058, W.M.; forest floor, Leith Stream bush, Dunedin, H 5582; in bush with Chiloscyphus billardieri, Bluff Hill, Southland, H 5146, K.W.A.

Stewart Island; Bank, Kaipipi Road, 9923, W.M.; Pegasus, H 6183, Dr J. Murray. The type was from Dusky Sound, coll. Menzies, 1791. Also from Tasmania.

Temnoma quadrifidum (Mitt.) Mitt. (Text-fig. 1, figs. 17-20) Jungermannia quadrifida Mitt., FI. Nov. Zel., 11, 128, 1854; Hook. f. Handh. N.Z. FI., 11, 504, 1867.

Temnoma quadrifida Mitt, in Handb. N.Z. FI., ii, Supp., 753, 1867. Blepharostoma quadrifidum (Mitt.) Pears. Journ. of Bot., 194, 1887; Steph., Spec. Hep., 3, 640, 1909.

Plants dioicous, variable, minute to small, inconspicuous, terrestrial, usually with other bryophytes. Stems simple or with 1 or 2 branches, one often from just a little below the perianth, nearly 1 cm long (Hooker gives 1 inch) barely 0.2 mm wide, cortical cells quadrate-rectangular to ca. 45/*, mostly less, x ca. 28-30 /* wide : branches about the same length. Leaves quadrifid to ca. small at the base of the stem, becoming progressively larger and more distant towards the perianth (on one stem), 0.6 mm long, as a rule not crowded, obliquely spreading from a narrow base, then curved so that the lobes are parallel with the stem, or with the lobes in the same plane as the basal portion, a little concave with transverse insertion, variable as to the width and armature of the lobes; base of lobes 0.15 mm wide (ca. 4-5 cells in minute forms), lobes lanceolate, tipped with a single cell or more rarely spinescent with ca. 3-4 single upper cells; margins usually entire but may be sparsely toothed. Cells of lobes 30 x 30-40/*, basal ca. 28-30/* x 60-70/*. Underleaves scarcely smaller and similar to the branch leaves with more slender lobes, at times coming close to stem leaves of T. quadripartitum. Floral leaves 1 mm long, reaching halfway or more up the perianth, 4-lobed, lobes toothed or spinescent both apically and laterally; floral underleaf similar in size, also 4-lobed with lobes mostly spinescent. Perianth terminal, ca. 1.5 mm, 3-gonous and sulcate when young, but becoming cylindrical with an open truncate mouth edged with rather stout setae, $ bracts in 2 or 3 rows, apical on stem or branch, not crowded, 4-lobed to 0.5 mm, but may be much shorter, conduplicate. Antheridia 0.1 mm with stalk ca. 30/* long.

Owing to differences in size, one cannot be precise as to leaf-measurements, but those given are mostly of the type.

Minute plants from a creek bed Mt Ruapehu, 352, G.0.K.5., with stems only 1.3 mm long excluding perianth, have no lateral toothing on the floral leaves at all. The perianth is 1 mm long, including the apical spines, which are 0.2 mm long, of ca, 5 cells*

Very small plants, H 5369, K.W.A., from amongst mosses and short growth on hillside facing south, sheltered by sparse manuka growth ca. 200 ft, west side of Lake Waihola, south of Dunedin, has strongly toothed floral leaves, dentatespinous, with stem leaves entire.

Throughout this species and in the type itself, one meets with leaves having uneven sinuses, the middle one somewhat deeper as is characteristic of T. palmatiloha. But this is not mentioned in the earlier descriptions, nor is it shown in Mitten’s drawing.

Other localities are—North Island: Damp shady clay bank, near Atiamuri, H 561, K.W.A.; 210, 211, 2852, from damp places on Kiwi Hills near Wairoa, E.A.H.; Wellington S. Berggren (approaching T. palmatum).

South Island: On earth with Meesia muelleri, etc., in damp ground in gully above Quartz Reef Hill, ca. 2,300 ft, near Naseby, Central Otago, H 5721; amongst Bartramia sp.

on damp patch in open ca. 2,000 ft, Naseby, H 5423, K.W.A.; near Clarendon, South of Dunedin, H 5383, K.W.A.

Owing to its small size this species is probably far more common than the few noted localities suggest.

The type was Colenso’s No. 1177, with other bryophytes, including Mitten’s Marchantia macropora.

Temnoma quadripartitum (Hook, senr.) Mitt. (Text-fig. 1, figs. 1-2) Jungermannia quadripartita Hook. senr. Musci Exot. 2, pi. 117, 1820. Jungermannia podophylla Angstr. Vet. Akad. p. 11, 1872. Blepharostoma quadripartitum Steph., Spec. Hep., iii, 639, 1909. Temnoma quadripartitum Mitt. Journ. Linn. Soc. Bot. 15, 65, 1876.

Plants monoicous, very small, terricolous with other bryophytes, light brown (the type blackish with green tips), moisture loving. Stems to 1 cm (in plants seen), filiform, simple or with few branches, cortical cells to 50/i, x 25/x, but usually shorter than that, ca. 35/x long, rhizoids purplish rose in tufts, but not common. Leaves remote, 3.5 mm, irregular in direction, sub-erect to spreading, a little concave, transversely inserted, quadripartite to /% or more rarely to i, from a base to 5 cells high, segments gradually tapering from a 3 or 4-celled base, the middle portion, of several pairs of twin cells, with a basal, lateral, ± horizontal tooth; all segments widely divaricating to an apical average width of 0.5 mm; cells everywhere smallish, ca. 20 jj. x 15/x, walls medium but not always clearly defined, trigones none. Underleaves a little smaller than, and similar to the leaves, variable, insertion reaching right across the stem, ca. 0.1 mm wide, then widening to 0.5 mm across the apex, 4-partite with a basal lateral tooth on each side, base to 5 cells deep, segments stiff, divaricating, median ones usually narrow, .tapering from a 3-celled base, lateral ones often from a 2-celled base, then 7-8 or less pairs of cells narrowed to 2-4 single cells, ending in a sharp point, cells of equal size throughout. Floral leaves 4-partite, larger than the cauline, specially the uppermost pair and underleaf, reaching 2 /i of the length of the perianth; segments narrow, with margins spinous toothed, and branched apical spines. Perianth 2 mm, strongly ventrally keeled, narrowed to the mouth, which is not open and truncate as in T. palmatum and T. quadrifidum, mouth dentate with teeth 2-3 cells long; cells of perianth to twice as long as those of the leaves. $ bracts in 3 pairs on a short branchlet near the base of a branch arising 0.25 mm below the perianth, much inflated, conduplicate, sub-erect, margins of lobes shortly dentate. Antheridia not seen.

The monoicous inflorescence was described from a specimen in the Riksmuseum, Sweden, from Nothofagus pumila forest, Sierra Sorondo, above Las Gotorras, Tierra del Fuego, in association with Riccardia florabunda, M 23, Rolf Santesson, 10/11/1940.

As Stephan! describes the species as dioicous, it is possibly polyoicous.

This species differs from T. quadrifida in the monoicous inflorescence, and in the more deeply divided leaves and the narrower leaf-segments. Occasional leaves at the base of the stem are much larger in every way, and might well belong to T. quadrifidum. They certainly show the relationship between the two species. In the original description W. J. Hooker writes: “There are smaller leaves mixed with the larger ones in this plant, which I would be disposed to consider as stipules (underleaves) if they were placed with any sort of regularity.” There is, of course, a regularity, not always apparent in the remoteness of both leaves and underleaves with their divaricating segments. The absence of spines in the cauline leaves will also separate the species from T. palmatum.

North Island: Mokai Patea, 5,000 ft, a spur of the Ruahines, in wet ground near a tarn, with other hepatics in tufts, 5950, A. P. Druce, February, 1951.

South Island: Amongst Conostomum sp. and Lepidozia sp. Arthur’s Pass, 3,000 ft, H 5441, Dr M. Holdsworth. Also from Kerguelen Island (Moseley) fide Mitten.

The type was from Staten Land, near Cape Horn, D. Menzies, 1787.

In the Riksmuseum, Sweden, is one small stem labelled “ Jung. int. Jung, tenacifolia Lord Auckland’s Group,” without doubt collected by Hooker, which Dr Arnell has identified as B. quadripartitum.

Temnoma palmatum (Lindb., ex Pears.) Schuster (Text-fig. 1, figs. 10-12) Blepharostoma palmatum Lindb. ex Pears. Journ. of Bot. 25, 193, 1887; Steph. Spec. Hep., 3, 638, 1909.

Temnoma palmatum (Lindb. ex Pears.) Schuster, Bryologist 62, 240, 1959.

Plants dioicous, small to medium, terrestrial or in rotting vegetation, rarely in a pure colony, usually with other bryophytes, lowly depressed or loosely scrambling, dingy to golden brown. Stems short and dense, 0.5 cm to 2 cm or even more when growing loosely and solitary, flexuous, little branched, 0.1-0.15 mm broad; cortical cells quadrate or shortly rectangular; rhizoids not seen, but described by Pearson as frequent, light brown or white, proceeding from the base of the underleaves. Leaves variable, setaceous, remote to closely imbricate, strongly succubous, almost longitudinally inserted, flabelliform with a base to 0.16 mm wide, 0.6 mm long including the spines, lobes very short to short, based on 2 to many cells, but always ending in a tapering spine ca. 6 single cells long, cells of spines to 90ju,, each cell narrowing slightly upwards, width of basal ones ca. 23ju,; upper cells of lamina ca. 25/x in diameter, basal cells quadrateoblong, elongated to as much as 40/x, may or may not be minutely asperate; trigones minute. Underleaves similar but shorter with a shorter lamina or leaf-base than the lateral leaves, and there may be 2 shortish lateral basal spines, oppositely placed. Floral leaves and underleaves imbricate and reaching well up the perianth, 4-lobed with lobes more pronounced than is usual in the stem leaves, and spinous both apically and laterally. Perianth short, 1.3 mm, cylindrical-obconic from a narrow base, no angles discernible though trigonous when young, with the 3rd angle ventral, mouth truncate, 0.7 mm wide, armed with spines similar to those of the leaves, to 6 cells long; setae long and much entangled, flattening to ca. 8 cells across, cells hyaline elongate to 1 mm long at times, or nearly so, apical cushion from which the valves originate, with 4-5 more or less hexagonal cells across, of varying size; capsule valves purplish wine, to 0.65 mm long x 0.2-0.3 mm wide, apparently 2 cells thick; spores most minute, scarcely 10ju, elaters curved and angled, ± long x 8-1 0/a diameter bispiralled. $ bracts in 4-6 pairs, conduplicate, with the margins 1-3 spinous; stems may be capitate with empty bracts. Antheridia single.

In some specimens there are short supernumerary spines along the leaf margins in between the four main spines, and lobe apices may in some cases be surmounted by more than one spine. Slender stems present in a gathering may bear leaves with very short leaf bases.

The relationship of this species to T. pulchellum is obvious, the cell-type and leaf-base of both species being the same, but T. palmatum is smaller and less spectacular with leaf-lobes shorter to non-existent, and leaves less imbricate and less regularly set. There are specimens referred to this species on account of their small size, but their leaf-lobes could belong to B. pulchellum.

The length of the setae or spines varies. It seems to be that on fertile stems which are shorter and with the leaves more crowded (in specimens seen), the spines are shorter than on more slender solitary stems.

North Island: Auckland, 6324, W. Dawbin; watercourse on Mt Hikurangi, 5,000 ft, with Riccardia sp. and Isotachis Lyallii, 10136, A.P.D.; earth under manuka, near Atiamuri, 6309; burnt log at edge of beech forest E. of Taupo, 2,500 ft, H 559, K.W.A.; Northern Ruahines, 6325, H.M.H.; Mokai Patea (branch range of Ruahines) high tussock meadow in boggy depression with sphagnum and Isotachis montana and Chiloscyphus Billardieri 5961, Kapakapanui with Lophocolea spinifera , 836, A.P.D.; N.W. Ruahines, 666, E.A.H.; with other bryophytes, 828, A.P.D.; Mt Egmont, 4 specimens, G.0.K.5.; rotting log, manuka scrub, Eastbourne, 2783, N. J. Butler; rotten log in bush near top of Mt Tauhara ca. 3,000 ft, H 1954; rotting log in bush at edge of Urewera County, H 1957; rotten log, Puaiti Bush, near Rotorua, H 1955; shady bank, E. of Taupo, H 1958, K.W.A.; Little Akatarawa Valley, on Schistochila Balfouriana, 11604, H.M.H.

South Island: Beech forest floor with other bryophytes, Arthur’s Pass, 6310, H. H. Allan, 6326, Mrs Vere; Sugar Loaf Bush, Gass, 20, W. R. Philipson; Bealey River, S. Berggren; Silver Peaks, near Dunedin, 17419, Herb. Bot. Div. D.5.1.R.; Dunedin, S. Berggren; shady bank, under heavy growth, S. of Dunedin, H 654; rotten log on floor of beech forest, Haast Pass, H 5948, K.W.A.; head of Lake Manapouri, H 973, G. Simpson; edge of small creek, 4,000 ft, head of Lake Tekapo (Canterbury), H 6086, D. Scott; Stillwater Base Gamp, Caswell Sound, V.D.Z.; Castle Hill, S. Berggren, 3865.

Stewart Island: Staircase Falls, Paterson’s Inlet, 436, W.M. Auckland Islands: Musgrave Peninsula, 2628, H. Guthrie Smith; “Cape Expedition”. Also examined, one from Mitcham, Victoria, in clay on walls of a disused quarry, G. I. Skewes, comm. J. H. Willis. Pearson also quotes Canberra Mountain, N.S.W. The type was from Otago Coll. Hector, No. 13, 1863, amongst Hypnum rutabulum.

Temnoma palmatilobum Hodgson comb. nov. (Text-fig. 1, figs. 3-9) Isotachis palmatiloba Hodgson, Review N.Z. Sp. Gen. Isotachis Rev. Bry. et Lich. T. 18, fasc. 1-2, 1949.

Plants dioicous, medium to robust/golden to dingy brown or greenish and somewhat shiny, moisture loving, in tufted or depressed mats, terrestrial, alpine. Stems simple or a little branched, matted together in tufts to 6 cm or shorter, 0.22 mm wide, cortical cells subquadrate to rectangular, to 30 x 100 [x. Leaves flabelliform, sub-remote to closely imbricate, transversely inserted from a base to 0.3 mm wide, widening to as much as 1.5 mm across the top, to 1 mm long, 2-lobed to the middle with lobes again divided to /3 or a little more of the leaf length; lobes triangular-lanceolate ca. 0.2 mm across the middle, acute or sub-acute, rarely obtuse, margins entire except for an occasional tooth on the outside margin of the ventral lobe. Cells ca. 25-30//,, smaller at the apex, elongated at the base ,to at least 60//,, cuticle minutely papillate. Underleaves very similar, smaller or quite equal in size, from a base of 0.3 mm widening to 1.2 mm. Floral leaves ca. 1.6 mm long x 1.5 mm across the top, may be more equally 4-lobed than the cauline, all segments shortly but strongly toothed on the margins. On the fertile stems the uppermost cauline leaves merge gradually into the floral leaves; floral underleaves 1.9 mm long x 1.1 mm broad at the apex, subequally 4-lobed, margins, lobes and leaf base a little toothed. Perianth 3.5 mm or longer from a

narrowed neck, ca. 0.7 mm wide, 3-gonous with a well-marked ventral keel, mouth lobed and toothed, teeth irregular, 1-3 cells long. All perianths examined were empty. $ bracts to 8 pairs counted on one stem, intercalary, contiguous on the stems, more removed on the branches, occasionally on one side of the branch only, bilobed, insertion transverse, conduplicate with the fold reaching to the middle sinus, thus forming a dorsal and ventral lobe, each being again bilobed, margins entire; antheridia not seen.

The sterile type from the Kaimanawas differs somewhat from the plants with perianths from Mt Egmont, in being shorter, tinged with green, a little glossy, leaves not, or scarcely imbricated along the stem, with some wide leaf-segments with slightly smaller cells. Moreover, in the clustered uppermost leaves the difference in depth of the middle sinus and the 2 lateral ones becomes less apparent.

This species is nearest to T. quadrifidum, the similarity to which is apparent in some of the branch leaves. The complete absence of spines separates it from T. pulchellum, T. palmatum, and T. corrugdtum. The subdivision of the two main lobes into two smaller ones is rare in the genus, and shows some kinship to the genus Lepicolea the type of which L. scolopendra (Hook, senr.) Dumor.t. is also a New Zealand plant and another of Menzies’ Dusky Sound collectings, 1792.

North Island: Stream bank, Kaimanawas, with Pohlia cruda (moss) 4,500 ft, 1062, A. P. Druce, No. 1329, Herb. E. A. Hodgson, type of Isotachis palmatiloba, Jan., 1947; dripping wet bank, Mt Egmont, ca. 4,000 ft, with Synphyogyna sp. and a dicranaceous moss, 1328, 1330, A.P.D., Dec., 1948.

South Island: Power Line Gully, Cass, North Canterbury, E 37, G.U., comm. Prof, W. R. Philipson, May, 1958.

Temnoma schusteri sp. nov.

Planta sterilis, parva, gracilis, scandens alias hepaticas. Caulis ad 1 cm longus, flexuosus, ramis paucis, cellulae corticis quadratae vel rectangulares ca. 40-50//,, parietibus crassis vitreis; rhizoidibus fasciculiformibus, truncatis vel irregulariter tuberosis, tuberis multicellulatis, rhizoidiferis, hyalinis. Folia remota, vel imbricata, suberecta, transverse inserta, obcuneata, vulgo tripartita, cum sinibus inaequalibus, raro quadripartita (parum bisbifida), segmentis divaricatis ca. 0.35-0.4 mm longis, longe acuminatis, base 2-4 cellulas lata, parietibus übique crassis, lamina ca. 0.15 mm longa, cellulae loborum quadratae ca. 30//,, laminae hexagonae ca. 35//,, luminae ca. 10//, forma irregulares. Amphigastria similia, basis aequans latitudinem caulinis, sinibus aequalibus. Cetera non visa.

Plants sterile, very small, brownish green or pale green in a cushion of Acromastigum Mooreanum. Stems to 1 cm, flexuous, little branched, brittle, barely 0.1 mm wide, thinner filiform stems bear remote bifid leaves; cortical cells quadrate to rectangular with very thick, transparent, brownish or pale green walls, to as much as 50/t long, though often shorter; rhizoids tufted, at the bases of underleaves or branches, colourless, or pale greenish brown, truncate or enlarged at the tip into a shapeless mass of cells which may again produce rhizoids. Leaves remote or imbricate, sub-erect, transverse, to 0.7 mm long, obcuneate, deeply to ca. f of the length, segments diverging ca. 0.35-0.4 mm long, longly tapering from a base of 2-4 cells, to a long series of single not elongated cells, one sinus a little deeper than the other. Cells fairly uniform in size, ca. 30-35//,, hexagonal in the lamina, and quadrate or sub-quadrate in the segments, walls everywhere thick and bulging irregularly. Underleaves similar and of equal size, the base reaching across the stem, sinuses equal.

Unfortunately this specimen is in very small quantity, and owing to the absence of perianths and androecia, one cannot be entirely sure that it is a

Temnoma. It differs from the other species of the genus in its type of cell with very thick walls, irregularly bulging into the interior portion of the cell. The usually tripartite leaves with the unequal length of the sinuses may suggest an evolvement from the bisbifid leaves, in which the third segment has failed to divide. Occasional 4-partite leaves are present on the stem. Temnoma Dusenii (Steph.) Schuster, from Patagonia, also has tripartite leaves with unequal sinuses, but the cells are quite different, elongated with thinnish walls, and the leaf segments are broader without the long series of single cells. The element of unevenness in the leaf-segments of this species together with the thick-walled cells suggests a near relationship with Lepicolea, but in .this genus there are zones of highly differentiated cells in the leaf laminae, and the plants themselves are very much larger.

Growing amongst Acromastigum Mooreanum, the dominant species in a mixture of Adelanthus falcatus, Chiloscyphus Billardieri, C. cymbaliferus, Lophocolea australis, Bazzania involuta, Pseudomarsupidium piliferum, Lepidozia sp. in subalpine forest, Table Hill, Stewart Island, 481, W. Martin, 5/2/47, subnumber 9355 Herb. E. A. H. ex No. 3137.

It is named for Dr R. M. Schuster, who has recently reinstated the genus Temnoma (1959).

Temnoma corrugatum (Steph.) Schuster (Text-fig. 2, figs. 23-25) Blepharostoma corrugatum Steph. Hedw. p. 315, 1893; Spec. Hep. 3, 63, 1909. Temnoma corrugatum Schuster Byrol. 62, 240, 1959.

Plants pale, greatly resembling Trichocolea lanata “ pulvinatim caespitosa ”. Stems simple or branched, semi-prostrate, to 3 cm long in plants seen (Stephani gives 6), 0.5 mm wide excluding spines, cortical cell cavities very small, rarely more than 3Op, long and 20/x broad. Leaves crowded, concave, obliquely spreading, broader than tall, 0.7 mm long, excluding spines, x 1.1 mm broad, from a base of 3.5 mm broad, 4-5-lobed, usually 4, divided to sinuses obtuse, margins recurved; lobes lanceolate, occasionally unequally bifid, base fairly broad, widely diverging, apex and margins armed with curved spines with 8 or less single narrow elongate cells, 50-70//, long x 28//, at the base, tapering to only 10//, wide at the apex; apical spines may be based on pairs of cells from a tapering lobe; cells divided by 2 walls wider apart at the ends, spines on the lobe margins 3-5 (Stephani gives 3), more or less in opposite pairs, arising at right angles to the lobe and ultimately curved. Cell cavities 20-25//, in the upper portion of the lobes, irregularly increasing in length to as much as 60//, at the base, the incrassate walls taking on the appearance of elongate and confluent trigones. Underleaves considerably smaller than the leaves and occasionally reduced to only 2 lobes, but otherwise fairly similar. No reproductive organs seen, nor are any described by Stephani, but Schuster is probably right in referring the species to Temnoma.

The pale colour, lanceolate lobes with curved apical and lateral spines, and the incrassate areolation of both leaves and stem cortex at once distinguish this species from T. pule helium. This species may be a connecting link between the genera Temnoma and Anoplostoma.

South Island: Revolver Gove, Preservation Inlet, coll. Dr H. H. Allan, Fiordland excursion, January to February, 1946, associated with Herherta alpina and Lepidozia spinosissima (syn. Lepicolea longifissa Steph.).

The type was from Stewart Island, coll. T. Kirk.

Anoplostoma gen. nov.

Basionym Blepharostoma Dumort. Recueil d’Observations sur le Jungermanniacees, 1835

Generitype A. vaginatum (Herz.) comb. nov. Blepharostoma vaginatum Herz.

Planta pallida virens vel rosacea, depressa, in rupibus; dioica. Gaulis ad 1.3 cm regulariter et dense pinnatus, cellulae corticis hexagonae, metientes ca. 30 /r, rhizoidibus purpureo-rosaceis, fasciculatis, truncatis vel apicibus ramosis; pinnis lateralibus, dense foliatis. Folia caulina 0.8 mm longa x 0.9 mm lata, asymmetrica, transverse deinde parum decurrentia, dorso contigua, lobis angustis, pinnatim ciliato-setigeris, setis hyalinis, uno-cellulatis; cellulae inaequales, 25 ad 40 fi, trigonis majusculis. Amphigastria caulina minora, ad medium 4-loba, lobis medianis majoribus, lateralibus, minoribus vel dentiformibus. Folia pinnarum confertissima, regularia, quamvis minora eis caulinum, laxe accumbentia quasi vaginata, triloba, lobis augustis, squarroso-patentibus, parce setulosis. Amphigastria ramulina 0.4 x 0.2 mm, lobis angustis, subintegris, apice armato, cellulis magnis, ca. 40 x 30j a uno latere connata. Folia floralia majora, 3-loba ad medium vel minus, marginibus setulosis. Amphigastrium florale subaequale, 4-fida. Perianthia in caule terminalia, uno (vel duobus) subflorale innovatione, magna, ad 5 mm longa, conica, supra triquetra, infra inflata, ore angustissimo, subintegro. Capsula immersa (ut videtur), firma ovalis, quadrivalva, sporae magnae, ad ca. 60/x, asperae, elateres 0.1-0.15 mm x vix 10/r. Antheridia solitaria in axillis foliorum ramorum apice caulis.

Anoplostoma gen. nov. sensu lato

Plants medium to robust, terrestrial or rupestral, pale or dingy green, roseate in M. vaginatus, gemmiferous in M. whiteleggei. Stems with lateral branching, pinnate in M. vaginatus, monopodially cymose in M. whiteleggei; rhizoids tufted, vinous-rose, clusters large or small. Cauline leaves remote to crowded, transverse to succubous, asymmetrically to sub-symmetrically 3 or 4-lobed, margins spinous-ciliate, cilia 1-celled. Cells with rounded or oval lumina with thick walls and trigones or ± hexagonal with small trigones. Underleaves 2- or 4-lobed, spinous-ciliate. Floral leaves similar to, but a little larger than the cauline. Perianth terminal with 1 or 2 subfloral innovations, large, triquetrous towards the apex maybe with smaller ridges, narrowed to a sub-entire mouth. $ bracts non-saccate.

Subgenus Anoplostoma

Sporae magnae ad 60jx diametro; ramis pinnatis, foliis rameis vaginalis, gemmis absentibus.

Typus Anoplostoma vaginatum (Herzog) comb. nov.

Subgenus Tricon a

Sporae multo minorcs, plus minusve diametro elaterum equantes, foliis superis forsitan gemmiferis, gemmis trigonis.

Typus Anoplostoma whiteleggei (C. & P.) comb. nov. Although the sporophyte characteristics of these 2 subgenera differ in that the ratio of the spore width to that of the elaters is 6:1 in subgenus Anoplostoma and 2:1 in subgenus Trigona, we do not think that this difference outweighs the resemblances of the gametophytes; the more or less similar leaves margined with

1-celled hyaline cilia, the non-erect habit, the rose-red rhizoids, the similar perianths and non-saccate $ bracts. Hence our retention of these 2 taxa in one genus.

The relationships of this new genus are complex, but its similarities to Ptilidium suggest that it be placed in the family Ptilidiaceae.

Anoplostoma vaginatum (Herzog) comb. nov. (Text-fig. 2, figs. 26-34.) Blepharostoma vaginatum Herz. Trans. Roy. Soc. N.Z., 65, 355, 1935.

Plants dioicous, medium in a dense, layered, low mat, rose-coloured, on rock. Stems to 1.3 cm, 0.2-0.27 mm broad, densely, laterally pinnately branched, all spreading from a central base, may be forked at the apex, each prong becoming again pinnately branched; heavily shaded plants may have elongated growths without pinnae with leaves like those of the pinnae or branches, rhizoids in clusters, rose-red, to 1.5 mm long, truncated or branched at the summit, clusters usually not numerous with few or many rhizoids; cortical cells smallish ca. 30 /*, hexagonal interior cells scarcely different in cross-section; pinnae or branches subequal, 2-4 mm long, 4 uppermost pairs becoming progressively shorter. Stem leaves arising from between the pinnae, upper portion pale, asymmetric, 0.8 mm long, 0.9 mm broad, ciliate-setulose, mainly transversely inserted, then seemingly a little and narrowly decurrent, 3-lobed, lobes unequal, margins ciliate-setulose; cells to 40/*, usually shorter ca. 25/*, different sizes intermixed, cavities ± oval, trigones medium, sometimes confluent, cells a little larger towards the base with smaller trigones, upper cells empty, lower often with chloroplasts around the margins; cilia one-celled to 0.2 mm long, with an obtuse point, long and short cilia may be mixed. Stem underleaves large 0.6 mm long x 1 mm broad, 4-fid to about the middle, lobes sparsely ciliated, the lateral lobes somewhat smaller. Branch leaves 0.65 mm long, tri-lobed to about halfway, lobes narrow, bases parallel with the axis, then squarrose curving upwards, imbricate, very regular, tri-lobed, to halfway lobes narrow with entire margins, or 1-2 pairs of ciliae oppositely placed. Branch underleaves | bilobed, ca. 0.4 mm long x 0.2 mm broad, connate on one side with leaf, lobes entire or with 1 or 2 cilia, cellular with large clear cells ca. 40 x 30/*, 2 cells broad at the apex, with base of 3 cells wide. Floral leaves in 2 rows, larger than the cauline to 1 mm or a little longer, 3-fid to halfway or less, the an deal lobe the largest, all ciliate-setulose; floral underleaves almost equal to the leaves in length, 4-fid, margins of lobes ciliate-setulose. Perianth terminal with one or two sub-floral innovations, large, to 5 mm long, fusiform (dry), pale, hyaline at the mouth, upper portion triquetrous and trisulcate, but with subsidiary ridges towards the apex so that the mouth is much narrowed, with short, broad acute lobes; 1 cell thick at the apex, 2 cells lower down, and 3-celled towards the base; calyptra near to /% the length of the perianth, apiculate and bearing old archegonia.

Sporophyte: stalk of capsule 4-6 cells across; capsule valves 1.5 mm long, 0.4-0.6 mm broad; the walls appear to be 3 cells thick, the outer walls dorso-ventrally compressed with dark cell walls which are nodularily thickened. Then there seems to be another tissue of cells with evenly thickened dark brown walls, unless this is just the inside walls of the outer layer, when the capsule would be only 2 cells thick. The inner tissue of cells of the capsule wall is rather tender and will part company with the others quite easily and break up into single cells if rubbed lightly between 2 slides on the microscope stage. These cells are turgidly oval

with rounded ends, transparent with transparent, curved (apparently spiral) thickenings as in the hyaline cells of Sphagnum. The actual cell walls are quite thin. Spores large to ca. 60/x, rough, elaters 0.1-0.15 mm x scarcely \ojx.

Young antheridia solitary, in axils of branch leaves near the apex of the stem.

This remarkable and beautiful species is at once recognizable even in the field by its rosy colour, densely pinnate stems and very hairy looking leaves margined with hyaline 1-celled curved cilia. The pinnate habit distinguishes it from A. whiteleggei, from which it also differs in the constantly 3-lobed asymmetric leaves, the vaginant branch leaves, and the usually bi-lobed branch underleaves. The perianths of the 2 species are similar, also the 1-celled marginal cilia.

So far this species is known only from the Central North Island Plateau, from the following stations: Sheltered corner of exposed bank on earth on side of track up Rainbow Mountain, near Rotorua, H 1965; banks of Rangitaiki River, on steep bank shade near Murupara, H 1963; rock face in gully, E. of Waiotapu V., ca. 1,700 ft, H 1964; “ Hongi’s Wall”, shaded rock face near source of Pongakawa River, Bay of Plenty, H 1960; 3 specimens from shady rock faces near Atiamuri, H 1966, H 1961, H 1959, H 4, rocky, shady face of gully near Taupo, all coll. K.W.A.; shady ignimbrite cliff, ca. 2,500 ft, Hauhangaroa Range, 7023, A.P.D.

The type was from steep rock face in gorge, near Atiamuri, K. W. Allison, 11/9/29, Hl6B.

Anoplostoma whiteleggei (Carr, et Pears.) comb. nov. (Text-fig. 2, figs. 35-39) Jungermannia whiteleggei Carr, et Pears. Proc. Linn. Soc. N.S.W. ser. 2,2, 1051, t. 32, 1887.

Blepharostoma whiteleggei (C. & P.) Steph. Spec. Hep., iii, 641, 1909.

Plants dioicous medium, terrestrial on poor light ground, often with species of Bryum, etc., in loose tufts, fragile, very light green tinted with brown or rose, may be gemmiferous. Stems semi-prostrate, primary arising close together, short ca. 3-10 mm but prolonged cymosely by 2-3 successive, may be thinner, stems or branches bringing the total length to 3 cm, the secondary stems arising laterally from a little below the apex of the preceding axis which then stops growing, sometimes 2 new thin stems arising near together; both primary and secondary stems end in dense clusters of leaves, width excluding leaves ca. 1.5 mm, cortical cells hard to discern but apparently linear-rectangular to more shortly quadrangular, interior cells numerous, in transverse section similar in shape to the leaf cells, rhizoids rare, but when present bright red-rose in tufts. Leaves variable, large, roughly symmetrical, densely clustered on the upper portion of the stem, strongly succubous, becoming deflexed on the lower portion of the stem and less crowded, ca. 2 mm long x 2.5 mm broad, reniform, mainly 4-lobed to about lobes with bulging marginal lobules sometimes overlapping on the adjacent lobe, or the marginal area folded back on the lobe itself, spinous-ciliate, sinuses obtuse, undulate, margins revolute; marginal cilia 0.1 to 0.3 mm long, 1-celled, hyaline, solitary or in clusters of 2-4, most often 3, with 2 of them and sometimes 3 arising from the same widened transparent base; when single they arise mostly from between 2 cells; short cilia are mixed with longer ones, straight or curved, the clusters arising from the apex of a lobe, lobule or a crenation; lateral single cilia in opposite pairs each set at right angles to the lobe. Underleaves smaller than the lateral leaves, spreading, 2lobed to halfway, with usually 2 smaller lateral lobes, ciliated as in xne leaves, connate on one side with the leaf or nearly so, the main sinus strongly recurved and more or less protruding. Cells fairly uniform throughout, ca. 30/x, a little larger at the leaf-base, but not elongated,

hexagonal with small trigones, walls medium to thin. Floral leaves 2.5 mm long, at the base of the perianth, scarcely larger than the cauline, 4-lobed (3 in one instance), with the 2 median lobes the larger, sinuses rounded with margins recurved, but not to the same extent as in the cauline; floral underleaf bilobed with a smaller lateral lobe on each side, not differentiated from the cauline. Perianth terminal on a primary stem, pale, in some cases tinted with rose, 3-5 mm long, oval, inflated, 2 mm wide across the middle and 2 cells thick, trisulcate and triquetrous in the main, tapering to a mouth on account of subsidiary plication, mouth ca. 0.7 mm wide, irregularly and shortly toothed, but not ciliate, whitish; cells similar to those of the leaves, clear in the upper portion, becoming opaque towards the base and somewhat more elongated. Seta of capsule ca. 3.5 mm long to ca. 0.2 mm wide; capsule valves dark, opaque, 1.1 mm long, 0.4-5 mm broad; spores of irregular shape, brownish, ca. 15ju, diameter; elaters 100-130ju, long x ca. l\\i wide. $ bracts in ca. 3 pairs at the stem apex, little altered from the ordinary leaves, ciliated but more sparsely, single antheridia at the base of each, ca. 0.2 mm long, slightly oval, shortly stalked, bracts not saccate.

Only one mature capsule was seen.

Gemmiferous plants erect, to 1.25 cm with 2-3 or perhaps more upper pairs of specialized smaller leaves having narrow-triangular lobes without cilia, but having instead, narrow, elongate, hyaline cells to 90 p. x 12/jl, not narrowed to a point. These cells are not at right angles to the leafmargin, but lie along it with apices projecting in the same plane. From these cells and it would seem from the inner ones as well, the gemmae are produced in quantities, and are frequently liberated and herd together on the slide. They are ca. 40ju, across, 3-cornered with concave sides, and 3-4 rounded cells well within the margins.

This also is a beautiful species, differing from A. vaginatum in the cymose, not pinnate, branching, in the bifid underleaves, and the more symmetrical 3-4-fid cauline leaves with shorter cilia. It resembles A. vaginatum in size, in the 1-celled cilia of the leaves, in the tufts of rose-red rhizoids, and in the large pale perianth with a plicate and shortly dentate mouth and relatively short floral leaves, and in the non-saccate $ bracts. In appearance A. whiteleggei is reminiscent of Temnoma corrugatum, but differs from this species in the single-celled cilia, and the constantly bifid underleaves, in the shorter stems and habit of growth, and in the wider and more closely leaved stems.

North Island: Under light fern on slope facing north, Waipoua Forest, H 770, East of Taupo ± 2,000 ft, 6314, hillside on earth amongst short fern, near Atiamuri, H 469, under manuka on dry ridge, Pongakawa River, Bay of Plenty, H 1972, under manuka on dry fiat, Tahorakuri Block, near Taupo, H 560, K.W.A.

Gemmiferous specimens were from dry slope facing south in tussock land, Kaingaroa State Forest Plantation, H 1971, East of Waiotapu Valley, Rotorua County, ca. 1,700 ft, H 1970, K.W.A.

The first collecting of this species in New Zealand, so far as is known, was from a hillside, on decaying tussock near Atiamuri, H3BO, 8/9/29, collected by K. W. Allison, as were all the subsequent gatherings.

In recent years specimens have been discovered near Melbourne by J. H. Willis, who has also received specimens from Mt Barker, in Western Australia.

The type was from New South Wales, collected by Whitelegge and Reader.

Bibliography

Carrington, 8., and Pearson, W. H., 1887. Hepaticae of N.S.W., Proc. of Linn. Soc. N.S.W., 11, p. 1035, Sydney.

Dumortier, B. G., 1835. Receuil d’ Obs. sur les Jungermanniacees, Tournay.

Fulford, M., and Taylor, J., 1960. Two new families of leafy Hepaticae; Vetaformaceae and Pseudolepicoleaceae from southern South America, Nova Hedwigia, 1, 405-422.

Herzog, Th., 1935. Descriptions of New Species of N.Z. Hepatics, Trans. Roy. Soc. of N.Z., 65, 355.

Hodgson, E. A., 1949. Review of N.Z. Species of the Genus Isotachis, Rev. Bry. et Lich., T. XVIII, fasc. 1-2, 25-31.

Hooker, Sir W. J., 1818. Musci Exotici, Tab. 114, London. Lindberg, S., 1887. In Pears. Journ. of Bot. 25, 193.

Mitten, W., 1864. FI. Nov. Zel., ii, 128. Journ. Linn. Soc. Bot., 15, p. 65.

Muller, K., 1948. Der System. Wert von Sporophytenmerk. bei den Beblatterten Lebermoosen Svensk Bot. Tid. 42, 1, Uppsala.

Schuster, R. M., 1958. Annotated Keys to the Orders and Gen. of Hep. north of Mexico, Bryologist 61, 1-66,

— 1959. Studies in Hep. 1 Temnoma, Bryologist, 62, 233.

Stephani, Fr., 1900. Beitrage zur Lebermoos-Flora Westpatagoniens und sxidlichen Chile, Bih. till K. Svens. Vet. Akad. Handl. Band 26, Avd. 111. N:r 6, Stockholm.

—1909. Spec. Hep. 111, 635.

E. A. Hodgson, Kiwi Valley, R.D., Wairoa, Hawkes Bay.

K. W. Allison, 9 Delta Street, Roslyn, Dunedin.

SUPPLEMENTARY NOTE

Since this paper went to press, we have been advised by Dr R. M. Schuster, at present of Otago University, that he has already described the new genus Anoplostoma under the two names, Herzogianthus Schuster and Chaetophyllopsis Schuster (1960-61).

Anoplostoma vaginatum (Herzog) Hodgson & Allison therefore constitutes a synonym of Herzogianthus vaginatus (Herzog) Schuster.

Whether or not Blepharostoma whiteleggei C. & P. represents still another genus Chaetophyllopsis whiteleggei (C. & P.) Schuster, must remain a matter of opinion, but in either case, Anoplostoma whiteleggei (C. & P.) Hodgson & Allison has been rendered invalid