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Volume 77, 1948-49
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New Zealand Hepaticae. (Liverworts). VI A Review Of The New Zealand Species
Of The Genus Frullania

[Read before the Hawke's Bay Branch, September 18, 1947; received by the Editor, November 18, 1947; issued separately, May, 1949.]

The genus Frullania, is very large, complicated, and highly specialised. Unlike the original genus Lejeunea, to which it is in some ways related, and which is now represented by 58 separate genera, the sub-divisions of Frullania, as initiated by Spruce, still remain sub-genera because, as Spruce says, “The outlying species of one group merge into those of the next, so that no limit is assignable, and (in my judgment) it is best to preserve the genus Frullania entire.”

Frullanias, on the whole, are not plants of the rain forest, but manuka scrub and kanuka are pre-eminently the favoured home of the majority of the species. Spruce remarks that in South America, some of the species have the habit of weeds, following in the tracks of man, and persisting on the edges of deserted clearings, on bushes and hedge-rows, and at the fords of streams; and he sees in this characteristic a possible explanation of their widespread distribution. To a small extent this is also the case in New Zealand, where I have noticed that F. squarrosula, F. pentapleura and F. patula are quite capable of taking up their abode on domesticated trees.

In the matter of perianths, Spruce also suggests that the rough perianth is an older form than the smooth one in which roughnesses and appendages may have become obsolete; while MacVicar regards perianth tubercles as possible gemmae, since similar tubercles may also appear on the leaves (in F. squarrosa and F. dilatata). The occasional appearance of minor ridges of varying lengths on the perianths of some species is very perplexing, though in species in which the perianths are normally ridged, such as F. cranialis, the rugulose appearance is constant.

The small ventral lobes or lobules are of two forms, the normal form characteristic of the species, and the explanate or evolute form. Both forms appear to be distinct even in their young stages. “So far as we are aware, there is not even an hypothesis as to why a shoot gives rise to saccate ventral lobes for a while, then suddenly changes to the explanate ones for a while.” *

Jubula Dum., the other genus in the family Frullaniaceae, as far as is known, is not represented in New Zealand.

Frullania Raddi

Frullania Raddi, Jung. Etrusc., Soc. ital. di Modena, 1820; Syn. Hep., 1845; Spruce, Hep. Amaz. et And., 1885; Schiffn., Nat. Pfl., i, 1893; Steph., Spec. Hep., iv, 1910; Verd., de Frull., i, 1928.

Plants small to large, creeping, matted or semi-tufted, red-brown to very dark or blackish, olive or pale green, sometimes tinted rose.

[Footnote] * Lois Clark and T. C. Frye in The Bryologist, 48, 1946, p. 56.

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Stems regularly to irregularly pinnately or bi-pinnately branched, branches laterally arising from axils of the leaves. Leaves incubous, alternate, shortly inserted, usually with a ligulate or rounded dorsal appendix, entire, sub-orbicular, ovate or oval, ventral lobe or lobule helmet-shaped, campanulate, or cylindrical-clavate opening downwards, or narrowed and extended downwards in a curve, on the side further from the stem, the opening ± obliquely facing the stem; sometimes explanate. Stylus sometimes present between the lobule and the stem. Stipules always present, variously shaped with rhizoids only rarely adhering. Cells small to medium, sometimes vittate at base of leaf, walls often sinuous. Invol. leaves larger, lobule always explanate, with or without lateral laciniae, invol. stipule may be connate on both sides with leaves. Perianth free, emergent or exserted, obovate to oblong, tri- or tetragonous (third angle postical), maybe with supplementary keels, margins waved, straight or toothed; inflated, cylindrical, sub-spherical or ventrally concave, rough or smooth, contracted at the apex to a tubular beak, irregularly ruptured by the exsertion of the capsule. Capsule on a short stalk, elaters unispiral, adhering to the capsule valves after dehiscence. ♂ inflorescence lateral, globose to oblong, bracts saccate, closely imbricate, sub-equally bi-lobed.

The genus is divided into at least seven sub-genera. Of these, certainly two are represented in New Zealand: Trachycolea Spruce and Diastoloba, Spruce. In the sub-genus Chonanthelia Spruce, the lobule is joined to the leaf by a wide carina, more or less parallel to the stem, thus making the dorsal side of the lobule much larger than the ventral. Stephani places his F. quinqueplicata (F. pentapleura) in this division (perhaps on account of the two ventral keels as given in Spruce's original diagnosis), and he suggests * that F. kirkii might also be placed here. However, perhaps all our species are just as suitably accommodated in Trachycolea.

Sub-genus Trachycolea Spruce
Subg. Trachycolea Spr., Hep. Amaz. et Andin., 31, 1885.
Subg. Galeiloba Steph., Spec. Hep., iv, 358, 1910.

Plants small to robust, lobule galeate, saccate or campanulate.

Sec. Vulgares Hodgson.

Plants medium to robust, lobules cucullate, cell-walls flexuous with nodulose trigones. Perianths broadly ventrally keeled or convex ventrally.

F. squarrosula (Tayl.) Syn. Hep.

F. pycnantha (Tayl.) Syn. Hep.

F. falciloba (Tayl.) Syn. Hep.

F. falciloba (Tayl.) Syn. Hep., var. setchellii (Pears) Hodgson comb. nov.

F. nicholsonii Hodg. nomen nov.

F. rostellata Mitt.

Sec. Acutilobae Verd.

Plants medium, rostrum of lobule acuminate or longly produced into a slender spine, straight or curved.

[Footnote] * Hedwigia, p. 145, 1894.

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F. hampeana Nees

F. subhampeana Hodgson

F. allanii Hodgson.

F. spinifera (Tayl.) Syn. Hep.

Sec. Campanulatae Hodgson

Plants small to medium, occasionally robust, lobules more or less campanulate, not less than 1 ½ times taller than broad.

F. incumbens Mitt.

F. anomola Hodgson

F. fugax (Tayl.) Syn. Hep.

F. fugax (Tayl.) Syn. Hep., var. media Hodgson

Sec. Brevilobulae Hodgson

Plants mostly small, lobules not taller than broad, stipules toothed or shouldered.

F. pentapleura (Tayl.) Syn. Hep.

F. cranialis (Tayl.) Syn. Hep.

F. solanderiana Col.

F. reptans Mitt.

Sec. Discilobae Hodgson

Plants usually robust, leaves mostly rotund, lobules and stipules not constant in size.

F. patula Mitt.

F. kirkii St.

F. subdeplanata St.

Sec. Irregulares Hodgson

Plants medium to extended, stipules variable in shape, with cells differing in shape from those of the leaves, lobules with mouth distant from the stem, asymmetric.

F. deplanata Mitt.

Key to Species.
1. Lobules acuminate to spiniferous (excluding the spiniferous form of F. patula) 2
Rostrum of lobules not spiniferous 5
2. Stipules not toothed, perianth cylindrical-inflated spinifera
Stipules not all toothed to strongly toothed, perianth ventrally keeled 3
3. Perianths rare, involucres numerous, stipules strongly toothed hampeana
Perianths frequent, stipules bluntly toothed to entire 4
4. Perianths beset with processes, well exserted, green when fresh subhampeana
Perianth smooth, plicate, shorter and broader, glaucous when fresh allanii
5. Lobules campanulate, taller than broad 6
Lobules not campanulate 9
6. Perianths tuberculate anomola
Perianths not tuberculate 7
7. Cell walls sinuous, leaves imbricate 8
Cell walls not sinuous, leaves not or slightly imbricate fugax
8. Lobules incumbent on the stem, perianths ventrally keeled incumbens
Lobules not incumbent on the stem, perianths dorsally and ventrally ribbed fugax var. media
9. Lobules as broad as tall, mainly symmetrical, plants small 10
Lobules longer than broad, asymmetrical, plants medium to robust 13
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10. Perianths many-ribbed 11
Perianths 2-ribbed ventrally 12
11. Leaves not or slightly imbricate, cell walls straight reptans
Leaves closely imbricate, cell, walls sinuous cranialis
12. Very pale, lobules inclined from the stem at varying angles to more than 90°, leaf margins usually gemmiferous, perianth may be narrowed to the apex solanderiana
Green to blackish, lobules more erect than inclined, perianth not narrowed at the apex (normally) pentapleura
13. Cell walls of leaves sinuous, perianth ventrally keeled or convex 14
Cell walls of leaves not sinuous, perianth ventrally concave (except in F. patula) 19
14. Leaves squarrose when moist 15
Leaves not squarrose when moist 16
15. Perianth smooth squarrosula
Perianth rough pycnantha
16. Perianth smooth, apices of leaves (strongly) and margins of stipules recurved falciloba
Perianth rough, stipules flat 17
17. Perianth strongly ventrally keeled, rostrum short setchellii
Perianth obovoid-spherical with edges obscure, rostrum long 18
18. Perianth with laminate, recurved laciniae, margins of stipules fairly even nicholsonii
Perianth laciniae ending in a series of single cells, erect, margins of stipules with a blunt tooth or uneven rostellata
19. Perianth not immersed, oblong, sides decurved, cells of stipules different from those of leaves (except sometimes in F. subdeplanata) 20
Perianth immersed or sub-immersed, cells constant throughout 21
20. Plants medium, lobules constant in size, stipules not reniform, though variable in shape deplanata
Plants robust, lobules large, stipules ♂ reniform subdeplanata
21. Always robust, upper part of perianth deflexed, invol. leaves not undulate and spreading lobules never spiniferous (as far as known) kirkii
Medium to robust, perianth sub-spherical, invol. leaves undulate and spreading, lobules sometimes spiniferous patula

In the descriptions which follow, the word “variable” is not used in a scientific sense, but merely to denote diversities; also, the word “stipules” is used in preference to “underleaves” or “amphigastria,” as being less cumbersome.

In denoting collectors, initials have been used as follows: K. W. A. (K. W. Allison); A. P. D. (A. P. Druce); A. L. H. (A. L. Hodgson); H. M. H. (H. M. Hodgson); E. A. H. (E. A. Hodgson); W. M. (W. Martin); L. B. M. (L. B. Moore); G. O. K. S. (G. O. K. Sainsbury); F. MacD. (F. MacDonald); V. D. Z. (V. D. Zotov).

Frullania squarrosula (Tayl.) Syn. Hep.

Jung. squarrosula Tayl. Lond. Journ. of Bot., 1845.

Frullania squarrosula. Syn. Hep., 412, 1844–1847; Fl. Nov. Zel., ii, 160; Handb. N.Z. Fl., ii, 535; St!, Spec. Hep., iv, 409, 1910.

Plants robust, in clumps, or cushions, or creeping, green to redor purplish-brown. Stems usually ca. 3–5 cm., much branched, branches short to long, alternate, conspicuous, leaving the stem at an acute angle. Leaves oval-rotund, basal portion amplexicaul, upper half squarrose, specially when moistened, or amplexicaul and imbricate with margins reflexed, sometimes scarious; undulate or plane, alternate, dorsal margins meeting each other at a right angle or less, with a zig-zag effect. Cells 20–30μ, marginal smaller, walls sinuous or slightly so, trigones and cells larger and clearer near the base with

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straighter walls. Lobules small, arched, mostly parallel with the stem, usually covered by the stipules. Stipules sub-rotund, sometimes keeled, sinus small. Invol. leaves complicated to examine, oval-rotund, lobule with a lateral segment, lanceolate or fancily shaped; stipule connate at the base with the involucral leaves, lingulate, with two variously shaped lateral segments, sinus straight; margins of all segments may be recurved. Perianth large, broadly keeled, or even inflated with an apical depression, smooth. ♂ stems bear capitate androecia ca. 1 mm. long to about twice that length with apices often recurved.

The squarrose leaves mark this species; though stems are often present on which the leaves are wholly amplexicaul, the latter at once become spreading on being moistened. This habit and characteristic appearance is shared by the smaller F. pycnantha, but this species has a rough perianth. There are also instances in which F. squarrosula comes very close to F. falciloba.

There is a variable species with a tuberculate perianth called F. squarrosa (R. B. N.) Dum., which is widely spread in the Southern Hemisphere. The relationship of F. squarrosula to this species is the subject of the following remark in the Flora Novae Zealandiae: E. squarrosula “has a very close resemblance to F. squarrosa, so that without perianths the plants might be confounded.” But a specimen very similar indeed to F. squarrosula, and having the necessary tuberculate perianths with markedly uneven edges, has been found at Conical Hill (Otago), leg. G. B. Rawlings, no. 17155 P. R. B. Herb.; and as smooth perianths, according to the authorities are not unknown on F. squarrosa, it may be that our New Zealand plant is more intimately connected with that species than is supposed. The New Zealand plant was the type of Taylor's F. squarrosula, and for the present it is convenient to let it remain as such, but I suspect that in due time it will be merged with F. squarrosa (R. B. N.) Dum.

North Island: On manuka, valley of Waipoua R., Rotorua, Taupo and Atiamuri districts, 15 specimens, K. W. A.; on kanuka, near Paeroa, A. L. H.; base of gum-tree by inlet of sea, Tauranga, 15 localities in Wairoa County, edge of L. Waikaremoana, all on bark and bases of trees and manuka, E. A. H.; Bay of Plenty, G. M. O'Malley; near Dannevirke, A. L. H.; Mt. Bruce (Masterton), N. K. Welsh; on laurel and oak-tree in park, on cabbage-tree stem, Masterton, W. M.; on bark, Mangaroa, 697, A. P. D.; on kanuka, north-west L. Wairarapa Rd., on manuka, river flat, Orongorongo Valley, H. M. H.; on oak-tree, Wallaceville, 1134, A. P. D.

South Island: Picton, also Marlborough, J. H. McMahon; Pelorus Bridge (Nelson), A. L. H.; on bark of beech, L. Roto-iti, 2,100 ft., G. O. K. S.; Travers Valley (Nelson), 2,300 ft., A. P. D.; Avalanche Peak (Arthur's Pass), H. M. H. and F. MacD.; forest margin, near Hokitika, W. M.; trees in bush, Maungatua Range (Dunedin), ca. 1,200 ft., K. W. A.

Stewart Is.: On stems of Senecio rotundifolia, W. M., 367.

Localities given in the Handbook are: B. of Islands, on lichens, J. D. Hooker; Port Nicholson and Titiokura, etc., Colenso; Wellington, Stephenson.

The following five species of Colenso's have been reduced to F. squarrosula by Stephani: F. cunninghamiana Col., F. pulvinata Col., F. viridis Col., F. ichthyostoma Col., F. rotundifolia Col,

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Frullania pycnantha (Tayl). Syn. Hep.

Jung. pycnontha Tayl. Lond. Journ. of Bot., 566, 1844.

Frullania pycnantha. Syn. Hep., 411, 1844–1847; Fl. Nov. Zel., ii, 160, 1855; Handb. N.Z. Fl., ii, 535, 1867; St., Spec. Hep., iv, 408, 1910; Rod., Tas. Bry., ii, 86, 1916.

Plants green or reddish, or purplish-brown to blackish, smaller than F. squarrosula with stems more intricately interwoven, otherwise similar, I think, in all respects, except for the perianths, which are covered with minute projections, 1–3 cells long. Also some of the branch lobules are occasionally acuminate.

F. laciniaeflora Pears. appears to be this species. The description of the leaves as “horizontally inserted, vaginate, imbricate, recurved or squarrose,” in conjunction with the rough perianth, can apply to only one New Zealand species: F. pycnantha.

Stephani (1892) refers three of Colenso's species to F. pycnantha: F. banksiana Col., F. diffusa Col. and F. echinella Col.

North Island: Southern descent of Mt. Te Wana, at the headwaters of the Waioeka R., B. Teague; Rotorua, Taupo and Atiamuri districts, 15 specimens, K. W. A.; Bay of Plenty, K. W. A., G. O. K. S.; Te Tetahi track, Mt. Tongariro, G. O. K. S.; on Nothofagus cliffortioides, National Park, L. B. M.; Tauhara Mt. (Taupo), G. O. Malley; common on trees in bushy gullies and on manuka, Wairoa County, Taihape, Palmerston North, 2 localities, E. A. H.; on kahikatea, Dannevirke, A. L. H.; Wanganui, 3 localities, K. W. A.; Southern Ruahines, 768, 741, Manawatu Gorge, 221, Kaipiti Is., A. P. D.; with Teloschistes on tree stem, Kopuaranga (Wairarapa), 385, W. M.; on kanuka, North-west L. Wairarapa Rd., H. M. H.; Orongorongo R., 7142, 7178, 7180, V. D. Z.

South Island: On manuka, Pelorus Bridge (Nelson), A. L. H.; log in bush, Glenledi (Milton), G. Simpson; tree trunk in bush, coastal ridge between Taieri and Akatore R., K. W. A.; Marlborough, J. H. McMahon.

The Handbook note is: North and South Islands, probably common, on lichens, etc., from B. of Islands to Thomson's Sound, Lyall.

Frullania falciloba (Tayl.). Syn. Hep.

Jung. falciloba Tayl. Lond. Journ. of Bot., 581, 1844.

Frull. falciloba. Syn. Hep., 423, 1844; Fl. Nov. Zel., ii, 160, 1855; Fl. Tas., 1860; Handb. N.Z. Fl., ii, 536, 1867; Rod., Tas. Bry., ii, 84.

Frull. colensoana St. Spec. Hep., iv, 1910; Pears, Univ. Cal. Pub. Bot., 10, 317, 1923.

Plants olive-green to golden- or red-brown to purplish black, medium to robust, in small dense, or large straggly cushions, on rock, trunks of trees in bush or open country and manuka scrub. Stems to 6 cm., but usually shorter, much branched, stems and main branches pinnate to bi-pinnate or sub-dichotomous. Leaves a little amplexicaul when dry, apices noticeably recurved, dorsal auricle triangular to rounded on the same stem, cells with flexuose walls; lobule arched, mostly parallel to the stem, half hidden by the stipules. Stipules large or medium, oval to obovate, may be a little auricled at the base, strongly keeled towards the base, margins recurved, apical notch small. Invol. leaves horizontally spreading, may be falcate, acute or sub-obtuse, about twice as long as broad, lobule with a lateral segment,

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stipule with 1–2 laterally toothed segments. Perianth large, with a broad obtuse ventral keel. ♂ stems simpler with perigonial bracts on short alternate branches.

A fine species, commonly met with, and well characterised by the strongly incurved leaf apices, together with gibbous stipules with margins also recurved.

The type was a Tasmanian plant, and Stephani does not record F. falciloba for New Zealand, but having seen specimens from Tasmania and one from North Queensland, 5976 N.Q. N.C. Herb., I am satisfied that Mitten was right in referring our New Zealand plants to that species.

F. colensoana St. is this species, and the type of F. falcata St. consists partly of this species and partly of F. deplanata.

North Island: Wairoa County, fairly common; E. A. H.; on limestone rock by sea, Mahia Peninsula, G. O. K. S.; on manuka and rock, near Atiamuri and E. of Taupo, 5 specimens, K. W. A.; bush near Rotorua, J. Lane; headwaters of Rangitikei Gorge, H. M. H.; cabbage tree, B. of Plenty, K. W. A.; on bark in bush, Taihape, E. A. H.; South Ruahines, 3, 730, 732, 781, Akatarawa V., 927, A. P. D.

South Island: Marlborough 2, Nelson, J. H. McMahon; Cobb Valley (Nelson), Pelorus Bridge (Nelson), A. L. H.; Travers V. (Nelson), A. P. D.; Mt. Arthur (Nelson), G. O. K. S.; Punch Bowl (Arthur's Pass), Foothills Torlesse Range, F. MacD.; Cass, Castle Hill on bark, Bealey Track (Arthur's Pass), Foothills Torlesse Range, Port Hills on rock, H. M. H.; near L. Harris, 17082, V. D. Z.; vicinity of Haast Pass, M. P. Matthews; near Mt. Cook, G. O. K. S.; near Fox Glacier, Mrs. Knight; on rock Mt. Watkin (Waikouaiti) 2, trees in forest near Dunedin, G. Simpson and J. S. Thomson; bark of trees, Mt. Cargill, 24647, P. R. B. Herb., in part, G. Simpson; Hitchen Hills (North Canterbury), 42077, P. R. B. Herb., hills above Scargill, A. J. Healy; on rock, Maungatua Range, ca. 2,500 ft., on rock on hillside near Berwick, rock on open bank of stream near Herbert, K. W. A.; Conical Hill (Otago), 17183, P. R. B. H., G. B. Rawlings; on rock, Alexandra, G. Simpson; on rock in gully, Horse Range, near Palmerston South, K. W. A.; Milford Track, F. MacD.

Stewart Island: 3 localities, W. M.

Var. setchellii (Pears.) Hodgson comb. nov.

Plants smaller and more tender, leaves approximate to a little imbricate, with the lobules reaching below the ventral margin, often showing from the dorsal view of the stem, stipules flat, perianths scabrous as in F. pycnantha, often faintly ribbed.

That this variety is really bound up with F. falciloba is shown by the following: Quite an otherwise normal plant of F. falciloba from beech bark, L. Roto-iti, Nelson, G. O. K. S., 1947, has the rough perianth. From the same locality are the smaller plants of the setchellii type, with smooth perianths; also from beech bark, Rangitikei V., A. P. D., 984, Jan., 1947. Again, a gathering of F. setchellii Pears., from shady rock on dry hillside, Ongaroto Rd., nr. Atiamuri, K. W. A., Feb., 1931, no. H267, has at least one smooth perianth, with numerous rough ones. Three specimens have smooth perianths, but the margins

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are lumpy; on manuka and on trees, Huka Falls (Taupo), Jan., 1940, H. M. H.; on upper branches of tawa tree, Maungapoike Falls (Wairoa), E. A. H., Dec., 1943.

Two gatherings with perianths apparently all rough are: On tree trunk in bush at edge of Urewera, K. W. A., 7/39, H547; on bark, “McKinnon's” bush, Wairoa, April, 1935, E. A. H. Sterile specimens may be scarcely distinguishable from F. falciloba, specially if the stipules are keeled. Localities for these are: Lower trunk of nikau palm in wet bush, Waipoua Forest, K. W. A.; on log with Dicranoloma sp., L. Waikare-iti, Morere bush, Tauruarau, on bark in bush Taihape Domain, E. A. H.; Waikaremoana, with Dicnemon calycinum, Mrs. H. Jeffreys; Kapakapanui, 895, A. P. D.; bush near Opepe, K. W. A.; Tauherenikau, V. D. Z.

The type, no. 213713, Herb. Univ. Cal., from bush near Waiotapu, is sterile, but even so, there can be no doubt but that the plant under discussion is the same as Setchell's, as it agrees with Pearson's drawing and description, and there is none other that the latter could refer to.

* Frullania nicholsonii Hodgson nomen nov.

F. berggrenii Nichols., The Bry., vol. 28, 1925; non St. (1924).

Plants olive or dark to purplish-brown, not common. Stems to 5 cm., irregularly branched. Leaves contiguous to imbricate, horizontally spreading, to about 1 mm. long, ovate-oblong, apex broadly rounded, a little recurved; smaller on branches, antical base with a ligulate appendix. Cells ca. 30μ, walls sinuous, trigones nodulose, basal vitta to ca. 40μ Lobule arched, cucullate, asymmetric, mouth narrowed, distant from the stem, and reaching below the leaf-margin. Stipules variable, but broadening from a transverse insertion and a narrow base, margins entire, sinus to ⅓ the length, segments acute. Invol. leaves oblong, apex obtuse, lobule as long, narrow-lanceolate, acute, margins recurved, margins of stipules unequally lacerate. Perianth pyriform, terminal on short branches, longly rostrate, beset with ligulate lamellate appendages with recurved apices.

The perianth of this species, “with somewhat the appearance of a miniature fir cone,” appears to be unique in the genus Frullania, but Lopholejeunea colensoi bears a similar one. In the absence of the perianth, the species can be mistaken for F. falciloba, the cells being similar; but the stipules of F. nicholsonii have flatter margins, and scarcely keeled, and the lobules are more distant from the stem, reaching well beyond the margin of the stipule.

On bark of tawa tree, Puaiti Bush, near Rotorua, K. W. A.; on bark of big tree, Ngamoko Track, Waikaremoana; on upper branches of tawa tree, with lichen, bush by Maungapoike Falls (Wairoa); Ohuka, on exposed log at edge of bush (probably), E. A. H.; Akatarawa V., almost certainly, A. P. D., 935.

Travers V., Nelson, with F. falciloba, A. P. D.

The type locality is given as Otira Gorge, South Island. 3346, Dr. S. Berggren.

[Footnote] * The name of F. berggrenii St. Spec. Hep. VI, 537, 1924, had already been bestowed on a South American plant (Bolivia) leg. Herzog.

Picture icon

Fig. 1—F. squarrosula. Fig. 2—F. pycnantha. Fig. 3—F. falciloba. Fig. 4—F. bergarenii. Fig. 5—F. rostellata. Fig. 6—F. hampeana. Fig. 7—F. subhampeana Fig. 8—F. allanii. Fig. 9—F. spinifera. Fig. 10—F. incumbens. Fig. 11—F. anomola. Fig. 12—F. fugax. Fig. 13—F. pentapleura. Fig. 14—F. cranialis. Fig. 16—F. solanderiana. Fig. 16—F. reptans. Fig. 17—F. patula. Fig. 18—F. kirkii. Fig. 19—F. subdeplanata. Fig. 20—F. deplanata. Fig. 21—F. rostrata. Fig. 22—F. aterrima. Fig. 23—F. ptychantha.

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Frullania rostellata Mitt.

F. rostellata Mitt. in Appen. to Handbook N.Z. Fl., if, 1867; St., Speo. Hep., iv, 410, 1910; Rod., Tas. Bry., ii, 86, 1916.

Plants green to red-brown, in extended patches, but not necessarily dense; may be gemmiferous. Stems to about 2 ½ cm., irregularly pinnate to bi-pinnate, branches of unequal length. Leaves oval-elliptic, a little imbricate, variable in size, but none longer than 0·8 mm. in specimens seen, flat and horizontally spreading (when moistened), ‘antical base extending across the stem, usually about as much as the stem is wide; lobules variable, swollen at the top, which may be contiguous to, or a little removed from the stem; gradually narrowing, dorsal side sometimes showing as wider than the ventral, diverging from the stem, i.e., the mouth is further from the stem than the middle top, mouth usually truncate; sometimes explanate. Cells fairly large, ca. 30μ, clear or opaque; trigones nodulose, walls flexuose, cell contents in an uneven ring. Stipules broadest across the middle, margins uneven, or crenate, or with lateral bulges, or bluntly toothed; sinus to about halfway, gaping, or with segments conniving somewhat. Invol. leaves in about 4 pairs on the short fruiting branches, increasing upwards and becoming more spreading; apices rounded, obtuse. Uppermost stipule elongate, broadest at the middle, may be toothed, bifid to nearly half its length, connate at the base with the involueral leaves. Perianth terminal on short fruiting branches, lateral on the main stem or primary branches, sub-pyriform, edges not pronounced, dorsal side flatter than the ventral, hispid with appendages, ca. 2–8 cells long, 1–3 cells broad at the base, extreme upper portion bare, rostrum conspicuous, 0 25 mm. long by 0·1 mm. broad.

The rounded, hispid, strongly beaked perianth is the main feature of distinction in this species. If perianths are absent, the broad, unevenly margined stipules are a good guide, together with the sinuous walled cells. In F. pycnantha the perianth is less rounded, with the appendages smaller and the leaves squarrose.

North Island: Little Barrier Is., on soft muddy rocks, Awarua Str., B. Molesworth; on kanuka in the Komata Str., near Paeroa; on kanuka on exposed rocks on coast near Colville. A. L. H.; Waitahanui R. Valley (B. of Plenty), K. W. A.; base of cabbage trees, on manuka, kanuka, light bush trees, often mixed with other Frullanias, Wairoa District, 15 specimens, E. A. H.; on rotten log, Whakamahia (Wai-roa), G. O. K. S.; Tauruarau, E. A. H.; Ohau-iti R., 7042, V. D. Z.; abundant on kanuka, north-west L. Wairarapa Rd., H. M. H.; coastal forest, Waikanae, 716, 721, Kapakapanui, 850, Kapiti Is., abundant, A. P. D.; on rocks near high-tide level, Titahi Bay (Wellington), W. M.; on Fuchsia bark, Wilton's Bush (Wellington), R. Mason.

South Island: Marlborough, J. H. McMahon; Mt. Stokes (Marl-borough), H. M. H.; Cobb Valley (Nelson), probably, A. L. Hodgson. North Island, on a Sticta, Knight, is given as the type locality.

Frullania hampeana Nees.

F. hampeana G. L., et N., Sun. Hep., 426; Fl. Nov. Zel., ii, 160, 1885; Handb. N.Z. Fl., ii, 536, 1867; Verd., Ann. Bry., Supp. Vol. i, 44, 1980.

Plants whitish or pale-green, often with tints of rose, reddish or even purplish-red, in flattish cushions on bases or trunks of trees or rock, or creeping on twigs. Stems irregularly pinnately to bi-pinnately

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branched. Leaves sub-remote to imbricate, ovate to oblong-ovate, apex rounded, obtuse, acute or even apiculate; flat or with margins slightly recurved. Lobule galeate with rostrum produced into a slender recurved spine, which may be as long as the lobule itself, lanceolate-acuminate when explanate. Cells to ca. 20μ, fairly clear, indistinctly 6-sided, trigones small or absent; cells of stipules irregular in shape with sinuous walls; stipules diversely shaped, usually taller than broad, insertion straight, sinus deep, gaping, margins irregularly 1–3 toothed, teeth sometimes spinous. Sterile inflorescences numerous, terminal on short branches. Invol. leaves in at least 2 pairs, acute to acuminate, lobules and stipule-segments acuminate-subulate, margins reflexed. Perianth oblong, flattish, rounded-truncate, 3-keeled on both faces, margins of keels with crenatures or hamate serrations, surface flecked with more or less tuberculate processes.

Only one perianth was observed, on a specimen from a shady rock near Atiamuri, K. W. A., 24/6/32.

Except that Verdoorn does not mention the perianth, our plants match his description very well (Verdoorn, 1930).

The many spinous-toothed stipules, together with the unusual colouring and ovate leaves with spiniferous lobules, make this species easy of recognition.

F. novae-zelandiae Col. is reported by Stephani to be F. hampeana Nees.

North Island: Edge of beech forest, Mt. Hikurangi, ca. 3,500ft., L. B. M.; 13 specimens from Atiamuri district, Puaiti Bush (Rotorua), K. W. A.; 9 specimens from Wairoa County, including bushedge at L. Waikaremoana and L. Waikare-iti, “Ngahere” (Puketitiri, H. B), beech bush, Tauruarau (Napier-Taihape Rd.), E. A. H.; near Ball's Clearing, Puketitiri, M. Brownlie; Birch Range (Kaweka), 3,000 ft., E. S. West; beech forest, Rangitikei V., 1024, 1042, Kapiti Island, A. P. D.; Tauhernikau, V. D. Z.; Wilton's Bush (Wellington), R. Mason.

South Island: Kenepuru (Marlborough), J. H. McMahon; east foot of Mt. Rubicon (Torlesse Range), H. M. H.; Abbott's Hill (Dunedin), 24701, P. R. B. Herb., G. Simpson; School Crk., Wakari (Dunedin), on scrub, Field Club, per E. Campbell; trunk of mahoe, near Herbert, K. W. A.; bush remnant, Otago Peninsula, on Aristotelia, Mt. Cargill, W. M.

Stewart Island, on stem of wineberry and Carpodetus, Oban, 491, 496, W. M.

Also found in Tahiti, New Caledonia, Java, Sumatra, Ceylon. Japan. The type was from Australia, in Herbarium Hampe.

Frullania subhampeana spec. nov.

Planta pallida, plerumque frugifer, dioica. Caulis ad ca. 2 cm., irregulariter pinnatim vel bi-pinnatim ramosus. Folia tenera, plerumque imbricata, ovata vel oblong-ovata ca. 0·5–0·7 mm. longa, sub-recte patula, obtusa. Lobuli galeati, rostro parvo vel longe attenuato. Amphigastria subrotunda, marginibus subintegerrimis, bifida, segmenta acuta. Perianthia ca. 2 mm. longa, obovato-oblonga, dorsale tri-sulcata, ventre convexa etiam tri-sulcata, plus minus laciniosa.

Plants pale-green to whitish, small, commonly fruiting. Stem to ca, 2 cm., irregularly pinnately to bi-pinnately branched. Leaves

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usually imbricate, ovate to oblong-ovate, margins a little reflexed or irregular in outline, may be finely papillose; upper portion of lobule swollen, rostrum elongate or short, recurved or pointing inwards towards the stem, often explanate. Stipules variable, but mostly suborbicular, lateral margins entire or with a blunt tooth which may be reduced to an angle or a sinuosity of the margin; sinus reaching to about ⅓ the height, gaping, segments acute, more diverging than conniving. Cells mostly clear with trigones confluent, cell-walls of lobules and stipules ♂ sinuous. Invol. leaves increasing in length upwards, acute, irregular in marginal outline, lobules and stipules usually less attenuated than in F. hampeana, ♂ lacerate. Perianth ca. 2 mm. × 1 mm. oblong or obovate-oblong, 3 plicate with a broad ventral keel usually with minor ridges, edges serrate with long teeth which may be hamate, crenate or irregularly dentate, surface flecked with processes, 1–5 cells long, base often more than 1 cell wide.

Distinguished from F. hampeana by the sub-entire stipules and the usual presence of perianths.

I would gladly have let this plant go as F. novae-zelandiae Col., but cannot be certain that F. novae-zelandiae is not a synonym of T. hampeana even although the “F. novae-zelandiae” in Stephani's herbarium (which is actually F. hampeana) is not Colenso's type. Colenso's description is vague, and Stephani's is a mixture of both species, so it seems safer to give this species another name, at any rate until the type in Colenso's handwriting is discovered.

Stephani lists F. monocera Tayl., from New Zealand also, but I do not think we have it here. A specimen from Rodway's herbarium, H 240a, shows it to have larger leaves than F. subhampeana, boldly toothed stipules, and a rough perianth partly hiden by the involucral leaves, which are toothed, all of which is in accordance with Taylor's description.

The following localities for F. subhampeana would show that it is not so widely distributed as F. hampeana: on bark of living tree, Puaiti Bush (Rotorua), K. W. A.; on bark in bush remnants round Wairoa, including Morere and Ruakituri, 10 specimens, E. A. H.; bark in bush gully, Panikau (Poverty Bay), G. O. K. S.; Tauherenikau, V. D. Z.; on bark of trees, Bledisloe Park, Palmerston North. E. A. H.; S. Ruahines (probably), Kapakapanui, 874, 876, Kapiti Island, A. P. D.; Wilton's Bush (Wellington), R. Mason.

Frullania allani sp. nov.

Planta viva glauco-viridis. Caulis ad 2 cm. longus, irregulariter ramosus. Folia imbricata, horizontaliter patentia, plana, oblongo-ovata vel elliptica, opaca; lobulis parvis, inferioribus galeatis, rostro recto vel curvato, longe attenuato, ceteris ac plurimis evolutis, subulatis, rectis antrorsum unispinosis. Amphigastria bifida, sub-rotunda integerrima vel lobis externe late obtuse unidentatis. Cellulae ca. 20μ. Foliis involucralibus triangulatis, acutis, integerrimis, lobulis magnis, triangulatis, marginibus planis vel recurvis. Perianthium late obovatum, 1 7mm. × 1·2 mm., leave, triplicato dorsaliter, carino tri-costato.

Plants medium, corticolous, bright bluish-green when fresh, fading to whitish in the packets. Stems rarely more than 2 cm., irregularly pinnately branched, Leaves broadly ovate to oblong-ovate

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flat, horizontally spreading; lobules mostly explanate, narrow, attenuated, divergent from the stem at an angle of about 45°, otherwise as in F. subhampeana. Invol. leaves increasing in length upwards, acute, ventral margins sinuous but entire; lobules large, triangular, from a wide keel, margins sinuous but practically entire, flat or reflexed. Perianth obovate, smooth, though lateral edges may be uneven, with 2 dorsal plicae, ventral keel with 3 ridges.

This species differs from F. subhampeana in the bluish-green colour when fresh, and in the shorter and broader perianth, which is ribbed and smooth. From F. monocera (Sec. Acutilobae, Tasmania), which it resembles in appearance, it differs in the less concealed, smooth perianth, in the invol. leaves being at most only sinuous instead of sharply toothed, and the cauline stipules mostly entire. F. acutiloba (Sec. Acutilobae, India and Ceylon) has a stem 5 cm. long, and the perianth has no supplementary keels.

A very similar plant incorrectly labelled F. hampeana, which I think might be this species, is from Toowoomba, Queensland, coll. C. Hartman.

On bark in bush remnant, “Rotonui,” Wairoa, June, 1931; 3 gatherings from bark in bush remnant, Kiwi Station, Feb., 1934, April, 1934, Easter, 1943; bush at Maungapoike Falls, Wairoa, Dec., 1931, E. A. Hodgson; tree-trunk in coastal forest, Terawhiti Hill, Nov., 1946, A. P. Druce.

Frullania spinifera (Tayl.) Syn. Hep.

F. spinifera Tayl., Land. Journ. of Bot., 407, 1846; Syn. Hep., Supp. 1847; Fl. Nov. Zel., ii, 161, 1855; Handb. N.Z. Fl., ii, 536, 1867; St., Spec. Sep., iv, 411, 1910.

Plants mostly olive-green, occasionally tinged reddish-brown, compact or straggly, usually corticolous. Stem to 4 cm., irregularly pinnately branched. Leaves imbricate to sub-remote, flat or with apices slightly incurved, ca. 0·7–1 mm. × 0·5–0·7 mm., ovate-elliptic, ovate or ovate-oblong, auricle rounded; lobule small galeate, acuminate, usually recurved, may be produced into a slender spine, often explanate, being then lanceolate, erect. Stipules medium to large, orbicular-cordate, auricled with an arched insertion, acutely bifid to about ⅓ the length, segments diverging or conniving. Cells rather opaque, arranged in rows, ca. 25μ, basal larger, may be brownish; walls thickish, usually straight, but may be sinuous, trigones usually present. Uppermost invol. leaves longest and narrowest, ca. 0·9 × 0·3 mm., lobule explanate, erect, reaching beyond the lobe by ¼–⅓ its length, bifid, inner segment short; connate with the stipule, which is as tall as the lobule, bifid to ca. ½, segments lanceolate-acuminate, armed, with a tooth on the outer side. Perianth tumid, sub-cylindric, sometimes with 3 shallow furrows, rostrum short, sub-immersed in an apical depression.

A distinct species, recognisable by the olive-green colour together with the small hooked lobules which are often explanate, and the sub-cylindric inflated perianth (when present). When packeted for a considerable time, it takes on a bluish mouldy appearance, due to the invasion of what looks like a filamentous alga.

North Island: Rawhiti (Bay of Islands), V. W. Lindauer; Waihau (Bay of Plenty), Mt. Tuhara (Taupo), G. M. O'Malley; Puaiti

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Bush (Rotorua), 3 gatherings, Oruanui (N. of Taupo), Roto-a-kui (B. of Taupo), between L. Rotoehu and the coast, 2 gatherings, Waihi (Bay of Plenty), Turakina (Wanganui), K. W. A.; numerous specimens from 7 stations in Wairoa district, E. A. H.; Poverty Bay, G. O. K. S.; Tarawera (Napier-Taupo Rd.), ca. 2,000ft., G. M. O'Malley; Esplanade Bush and Bledisloe Park (Palmerston North), abundant, E. A. H.; Wairongomai R., 7223, 7235, Ohau-iti R., 704], 7025, V. D. Z.; on kanuka, north-west L. Wairarapa Rd., H. M. H.; Terawhiti Hill, 821, 823, A. P. D.

South Island: Ashley R. Gorge, Banks Peninsula, Kennedy's Bush (Christchurch), Castle Hill, H. M. H.; Marlborough, G. O. K. S.; on tree trunk, bush near Dunedin, on tree branch in open bush in gully near Herbert (Oamaru), H822, K. W. A.; on bark in forest remnant, Otago Peninsula, W. M.

Auckland, Sinclair, Colenso on trees in forest, Tarawera.

Frullania incumbens Mitt.

F. incumbens Mitt., Fl. Nov. Zel., ii, 162; Handb. N. Z. Fl., ii, 537, 1867; Spec. Hep., iv, 406, 1910.

Plants small to robust, variable, straggly or compact. Stems may be extended to 5 cm., branches alternate, pinnate to bi-pinnate, the upper ones becoming more distant and shorter. Leaves contiguous to closely imbricate, usually spreading, concave, apex incurved, oblongelliptic, size varying in proportion to that of the stem. Lobule campanulate, about 1 ½ to 2 as long as broad, oblique, the upper portion usually obscuring the stem and stipules, mouth dilated. Stipules ♂ obovate to obcuneate, broader in robust specimens, specially on the main stems, often with an obtuse shoulder angle, sinus narrow. Cells ca. 18μ, walls flexuous, trigones nodulose, basal larger and clearer, to ca. 35μ. Invol. leaves much incurved, lobule with a lateral tooth, stipule ovate, bifid, segments ovate, acute. Perianth triquetrous, dorsal face usually convex, but may be concave, smooth, may be plicate at the mouth. ♂ branches sinuous with a median furrow, bracts in pairs to about 12, tightly packed.

The large, oblique, campanulate lobules and the smooth perianth ventrally keeled, are the distinguishing marks of this species, but it is very variable in size, the Arthur's Pass and North Canterbury specimens being very large.

North Island: Mostly on manuka, Rotorua, Taupo and Atiamuri districts, 26 specimens, K. W. A.; on bark of young beech, Whakapapa (Mt. Ruapehu), G. O. K. S.; on trees near Huka Falls (Taupo), H. M. H.; beech forest, Rangitikei V., 3,000ft., A. P. D., 1037; on manuka and bark of bush trees, Wairoa County, including Waikaremoana, 15 gatherings, on bark in open bush, Taihape, E. A. H.; foothills near Dannevirke, A. L. H.; Orua Valley, on manuka trees, river flat, Orongorongo V., H. M. H.

South Island: Mostly on beech bark, Arthur's Pass (Punch Bowl, Bealey Track, Avalanche Peak), H. M. H., F. Macd., W. M., 88 in part; tree trunk, Kelly's Hill (Otira), 379, W. M.; on beech bark, L. Roto-iti, 2,100 ft., G. O. K S.; Cass, Castle Hill, H. M. H.; Upper Waimakariri, M. Berry.

The type was collected by Colenso, and from the fragment seen, appears to have been a small plant.

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Frullania anomola sp. nov.

Planta dioica corticola, repens, olivacea vel rufescens. Caulis ad 2cm., irregulariter ramosus, rami saepe breves. Folia caulina ca. 1·1 × 1 mm., sub-remota, late cordata, ramulina ovalia, contigua vel imbricata, apicibus parum incurvis; lobulis campanulatis, caulinis remotis, erectis, ramulinis plus minusve confertis, saepe juxtapositis, incumbentibus. Cellulae ca. 25μ., parietibus multo sinuosis, trigonis majusculis vel maximis, saepe confluentibus. Amphigastria anguste obcuneata, sublobata in summis later ibus, bifida fere ad medium, segmentis divergentibus obtusis. Perianthia pyriformia vel oblonga, tuberculata, ventre late unicarinata, marginibus lateralibus tuberculosis, terminalia vel in ramulis lateralibus brevibus; foliis involucralibus oblongis, obtusis, lobulis latis, acuminatis, unilaciniatis; amphigastriis angustate oblongis. Mascula desunt.

Plants corticolous, creeping, olive-green to red-brown in the same gathering. Stems to ca. 2 cm., very irregularly pinnately to bipinnately branched, primary branches often short; terminally fruiting stems and branches branched a short distance below the inflorescence, usually on both sides. Cauline leaves sub-remote to contiguous, or a little imbricate, cordate, branch leaves smaller, broadly oval, more crowded with apices incurved; lobules ca. 0·4 mm. × 0·25 mm., campanulate, erect on the main stem, but may be oblique on the branches as in F. incumbens, when the mouth may be narrowed; cells ca. 25μ, markedly sinuous, trigones inedium to large, sometimes confluent. Stipules obcuneate from a narrow elongate base, shouldered or with a shoulder lobe, upper margins may be waved or crenate, bifid to nearly ½, segments obtuse diverging. Invol. leaves oval, lobule broad, acuminate with a lateral segment, margins reflexed; stipules narrow-oblong, bifid to about ⅓, uppermost stipule broader and unevenly bifid (in one case). Perianth pyriform to oblong from a smooth narrow base, tuberculate, broadly ventrally keeled, sometimes faintly ridged, edges bluntly toothed.

This species is nearest to F. incumbens, of which it might possibly be a parallel rough-perianthed form. But apart from the perianth, the lax habit, with scarcely imbricated leaves, exposing the black stem, give it a decidedly different appearance. So involved can the classification of Frullania become, that one could imagine that, with its tuberculate perianth, non-squarrose leaves and campanulate lobules, this species might be the Indo-Malayan plant Frullania squarrosa (R. B. N.) Dum., var. planescens Verd., forma campanuloides Verd.

On bark of Olearia sp. bushes, 3,500ft., Tararuas, H. M. H., Jan., 1947.

Frullania fugax (Tayl.), Syn. Hep.

Jung. fugax Tayl., Lond. Journ. of Bot., 1845; Frullania fugax, Syn. Hep., 445, 1844–47; Fl. Nov. Zel., ii, 161, 1855; Handb. N.Z. Fl., ii, 537, 1867; Spec. Hep., iv, 1910.

Plants small, creeping, reddish-brown, often gemmiferous, rarely fruiting. Stems to ca. 2 cm., irregularly pinnately branched, stems and branches often bare of leaves with some stipules showing for parts of their length. Leaves fugacious, orbicular-oval, 0·4–0·5mm., contiguous or sub-imbricate, diverging, the antical base covering the stem, auricle rounded; lobules large for the size of the leaf, campanulate

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as in F. incumbens, 1 ½ to 2 as long as broad, rarely with a constricted mouth, erect or a little oblique. Stipules obovate, may be rather abruptly widened near the top with an obtuse shoulder angle; sinus usually gaping. Invol. leaves about twice as long as broad, apex rounded, lobule acute to acuminate, with a small lateral segment, stipule with segments and margins of segments noticeably recurved. Perianth with 2–3 dorsal ribs and 2–3 main ventral ribs; rostrum long.

In appearance and cell-structure, this species resembles F. pentapleura, but differs in the longer lobule and caducous leaves, though these are not unknown in both F. pentapleura and F. reptans. The following observation of Pearson's, regarding a plant of Setchell's, collected in bush near Waiotapu, is very much to the point: “In wetting the plant for examination the leaves nearly all fall off,… The plant has a reddish colour, the lobule very large and underleaves small. I found numerous involucres but no perianths. Gemmae are very abundant on the margins of the leaves.” The Handbook also has a note to the effect that the perianths are not fully formed.

North Island: Great Barrier Is., T. Kirk, 6134, P. R. B. Herb.; bark of trees, C. Brett, V. W. Lindauer; on manuka, valley of Waipoua R., K. W. A.; on kauri, Moehau, L. B. M.; on trees in Komata Stream (Paeroa), A. L. H.; on rotten leg in shade near Atiamuri, on manuka, E. of Waiotapu V. (light green), on tree bark, Puaiti Bush, on pohutukawas, Ohope Beach, H352, K. W. A.; Wairoa County including Morere and Waikaremoana, 7 specimens, on beech bark, Tauruarau (Napier-Taihape Rd.), abundant, E. A. H.; beech forest, Rangitikei V. (Kaimanawas), 3,000ft., 982, 1000, Mangaroa, 699, Upper Hutt, 913, 917, Kapiti Is., A. P. D.; on manuka trees, river flat, Orongorongo V., H. M. H.

South Island: Pelorus Bridge (Nelson), A. L. H.: beech bark, L. Roto-iti, 2,100 ft., G. O. K. S.

Var. media Hodgson

Planta compactior, ramosa irregulariter, rami et ramuli breviores, apicibis sub-incurvibus. Folia imbricata, incurva, majora. Parietes cellularum sinuosi. Perianthia oblonga, oblongo-ovata, vel anguste ovata, pluriplicata, carinulis sinuosis et crenulatis. Bracteis ad 10-jugis, sub-circinatis (H510, K. W. A.).

This so-called variety appears to be intermediate between, and equally related to, both F. fugax and F. incumbens. But for classification purposes, I have made it a variety of F. fugax, on account of the ribbed perianth and the erect somewhat narrow lobule. It resembles F. incumbens in the shorter branches and imbricate leaves, and the sinuous walls of the cells. Even so, it is difficult to draw a hard-and-fast line between F. fugax and the var. media, and between the var. and F. incumbens. For instance, a second specimen from Pelorus Bridge, Nelson, has all the habit of F. fugax, with the sinuous cellwalls of var. media.

North Island: On manuka bark, Rotorua, Taupo and Atiamuri districts, 15 specimens, K. W. A.; on manuka, Ohuka (Wairoa), on manuka, “McKinnon's” Bush (Wairoa), beech Bush, Tauruarau, on manuka, Otupae, E. A. H.; Tauherenikau (may be F. incumbens), V. D. Z.; Akatarawa V., 928, A. P. D.: on manuka trees, river flat, Orongorongo V., H. M. H.

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South Island: Near Cass University Hut, H. M. H.; Happy Valley (Springfield), F. MacD.; on beech bark, L. Roto-iti, 2,100ft. (leaves caducous as in F. fugax), Governor's Bush (Mt. Cook), G. O. K. S.; Mt. Watkin (Waikouaiti), G. Simpson and J. S. Thomson.

The type of F. fugax was on Parmelia reticulata, New Zealand.

Frullania pentapleura Tayl.

F. pentapleura Tayl., Nov. Hep., Land. Journ. of Bot., 1846; Syn. Hep., 775, 1846; Fl. Nov. Zel., ii, 162; Handb. N.Z. Fl., ii, 537, 1867; St., Spec. Hep., iv, 414, 1910; Rod., Tas. Bry., ii, 87, 1916.

F. quinqueplicata St., Spec. Hep., iv, 356, 1910.

Plants minute to small, often gemmiferous, pale olive-green to reddish-brown or purplish-black, in thin patches on bark of manuka or in light bush, or on rocks, specially on the sea coast. Stems to 2 cm., irregularly pinnate to bi-pinnate, branches of varying length. Leaves small, larger on the main stem, ca. 0·5 mm., diverging, overlapping the stem, a little concave, oval-elliptic, remote to imbricate, may be caducous as in F. fugax; lobules variable, often explanate or semi-explanate, a little longer than broad, parallel with the stem, or inclined with the upper portion further away, lips not always parallel or at right angles to the stem. Stipules cuneate or sub-orbicular, bi-fid, segments sub-acute to obtuse, often diverging, mostly with a lateral angle or tooth. Cells ca. 20μ, usually rounded-quadrate, clear or opaque arrangement of cell contents may give the impression of sinuous walls; trigones small or absent. Invol. leaves oval with a wide keel and large lobule which has a small lateral segment. Perianth normally with 2 dorsal and 2 ventral keels, which are often wavy, edges entire or irreguarly crenate, apex rounded-truncate; ♂ spikes with 2–3 pairs of bracts.

The original of this species was from Australia, and Mitten recognised the species in our New Zealand plants. Having seen Taylor's type, I can see no reason why Stephani should disagree with this and call our New Zealand plants F. quinqueplicata, the type of which I have also seen.

North Island: Waiheke Is., Maria Is., Crusoe Is., on pohutukawa. Matiatia Bay on Opuntia stem, L. B. M.; Little Barrier Is. on cliff beach. B. E. Molesworth; Bay of Islands on foreshore rocks, shallow soil on rock, Waipoua R., K. W. A.; on bark, Ohope, H. M. H.; Raukokore R., G. O'Malley; Mt. Maunganui on pohutukawa trees, rocks on Blowhole near Mt. Maunganui (Tauranga), base of willows by inlet of sea, Tauranga, very plentiful in Wairoa County on manuka and bark in light bush, edge of L. Waikaremoana, Run 38 on rocks by Mohaka R., Havelock North at base of blue-gum, Tauruarau (Napier-Taihape Rd.) on kanuka, Otupae and Erewhon on bark and ground, on bark, Esplanade Bush, Palmerston North, E. A. H.; Dannevirke on bark, A. L. H.; bank of Rangitaiki R., K. W. A.; South Ruahines, 729, Akatarawa V., 938, robust with fertile stem, leaves imbricate as in F. fugax var. media, 936, Kohekohe, dominant coastal forest near Plimmerton, A. P. D.

South Island: Marlborough, J. H. McMahon; Lamb Hill, near Hindon, 24708, P. R. B. Herb., G. Simpson; on bare rock near Qutram, K. W. A.

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Frullania cranialis (Tayl.), Syn. Hep.

Jung, cranialis Tayl., Lond. Journ. of Bot., 86, 1845.

Frullania cranialis, Syn. Hep., 419; Handb. N.Z. Fl., ii, 536, 1867; St., Spec. Hep., Iv, 415, 1910; Rod., Tas. Bry., ii, 86, 1916.

Plants olive-green to red-brown, intricately matted. Stems much branched, branches of varying lengths. Leaves to 0·6 mm., closely imbricate, oval, apices incurved, loosely imbricate on outlying branches; cells irregular, ca. 20μ, increasing in size towards the base to as much as 40μ, basal hyaline, walls all sinuous. Lobules about as tall as broad, parallel with the stem, upper part rounded, sides parallel; the ventral basal portion appears sometimes to be appressed to the dorsal, thus causing the lips to be not parallel. Invol. leaves broad, lobule with a lateral segment, margins recurved. Perianths numerous, terminal on branches, obovate-oblong, strongly ribbed, with about 3 ribs on each face, occasionally only 2, usually waved, with margins rugulose. ♂ bracts in spikes consisting of 3–10 pairs.

The perianths of this species are very convincingly ribbed, with all cell-walls strongly sinuous as in the Australian type. It is normally a more robust plant than F. reptans, with leaves contiguous to imbricate and different cells. It is closely related to F. fugax var. media, from which it differs in the short lobule.

The plant on the Kew sheet, from Canterbury, leg. Haast, 1862, is labelled F. cranialis, but it has the long lobule of the fugax group.

A robust creeping ♂ specimen from Cobb Valley, Nelson (A. L. Hodgson), has loosely imbricate leaves and cells nearer to those of F. reptans. The ♂ branches are both short and long on the same stem.

H243, Herb. Rodway, determined as F. cranialis by Stephani, is either F. fugax or F. incumbens (perianths would decide). Neither of these 2 species is recorded for Tasmania by Rodway.

I would venture to suggest that F. macraeana, from Macrae's Island, near Wilson's Promontory, coll. F. von Muller, is F. cranialis.

South Island: Mt. Cheeseman (Canterbury), Mingha Valley near Arthur's Pass, F. MacD.; Governor's Bush, Mt. Cook, G. O. K. S.; Wilkin R. to L. Wanaka, G. Simpson; Castle Hill; H. M. H.

The type was from King George Sound, coll. Cunningham.

Frullania solanderiana Col.

F. solanderiana, Col., Trans. Roy. Soc. N.Z., 21, p. 75, 1888.

F. decurviloba St., Spec. Hep., iv, 409, 1910.

Plants small, pale, almost always gemmiferous, on twigs, associated with a blue Metzgeria sp., bark, rock. Stems intricate, tripinnate. Leaves very pale, very small, 0·3–0·5 mm., the larger ones on the main stems, broadly oval, remote to a little imbricate, “sub-margined,” margins usually gemmiferous. Lobule often a darker colour than the leaves, similar to those of F. pentapleura, but many of them more patent to horizontal, projecting beyond the margin of the leaf. Cells “clear of various shapes, subquadrilateral and irregularly angular, walls thick and double with minute cellules in them.” Stipules and invol. leaves as in F. pentapleura. Perianth a little narrowed to the apex, dorsal surface 1·2 ribbed, ventral keel with minor ridges, often waved and crenulate. ♂ branches as in F. pentapleura.

In a specimen growing on Aristotelia fruticosa twigs, near Atiamuri, H320, K. W. A., some of the marginal gemmae appear to have developed into minute, imperfect stems with leaves.

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The uneven leaf-margins, with the lobules often, projecting, give the plant a curious appearance.

Nearest to F. pentapleura, from which it differs in the usually pale colour, the more oblique lobules, the somewhat narrowed perianth, and in being more consistently gemmiparous.

The type stem, A1346, Frullania solanderiana, 1890, in Colenso's handwriting, on. a sheet in Kew Herbárium, has been re-named F. cranialis Tayl. by Stephani, while a piece of the same stem, A1346, is in Stephani's Herbarium, labelled F. decurvifolia St., sp. nov.

North Island: On twigs, Hopuruahine (Urewera), G. O. K. S.; on Fuchsia excorticata, Heipipi (Urewera), L. B. M.; on manuka twigs, on shady rock face in gully, on trunk of pine, Kaingaroa Plains, on lianes and dead fern fronds, on manuka, on cabbage tree stems, all east of Waiotapu Valley (Rotorua region), K. W. A.

Frullania reptans Mitt.

F. reptans Mitt., Fl. Nov. Zel., ii, 161; Handb. N.Z. Fl., ii, 536; St., Spec. Hep., iv, 1911; Pears., Kew. Bull, Misc. Inf., 2, p. 69, 1924.

This species has the habit, leaves, leaf-cells and lobules of F. pentapleura, but the perianth is pluriplicate and ♂ elongate as in F. cranialis. It is apparently rare, and not a very satisfactory species, but stich specimens do exist, and for the present cannot be explained away, unless as a small form of F. cranialis.

Such a one, both ♂ and ♂, is from a shady rock face, E. of Taupo, ± 2,000 ft., K. W. A., 12/11/34. The ♂ branches have up to at least 10 pairs of bracts, as in Pearson's description of the type. Another, more tufted than Mitten's type, is from the base of a tree, Run 38, by Mohaka River, E. A. H., April, 1938. One from manuka bark, side of gorge, “Raumai,” Wairoa, is clearly F. pentapleura with a modified perianth.

A sterile blackish gathering, widely extended, from wet ground near stream, Hyde, Otago Central, doubtless belongs here, as the lobules appear to be all erect. In F. pentapleura a proportion of them are usually a little oblique; coll. G. Simpson and J. S. Thomson, 25/9/35.

“East Coast on bark of yellow kowhai, Colenso; Auckland, Sinclair.”

Also reported from Tasmania.

Var. integristipula Nicholson

“3400 from Bay of Islands (S. Berggren) agrees in general characters with F. reptans Mitt., but differs from the type in the large entire underleaves. I propose to call it var. integristipula of that species.”

I have not yet seen any New Zealand specimen of Frullania with entire stipules.

Frullania patula Mitt.

F. patula Mitt., Fl. Nov. Zel., ii, 159; Handb. N.Z. Fl., ii, 535; St., Spec. Hep., lv, 413, 1910.

Plants usually robust, variable, creeping on twigs, or in dense mats on bark, sometimes widely extended, usually dull-olive-green, sometimes a little coloured. Stems to about 3 cm., branches numerous, short, mostly alternately pinnate. Leaves round, rarely oval, dorsally

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overlapping the stem, sometimes divergent, some semi-amplexicaul, or if crowded, rising up from the stem, when the margins may be undulate or scarious; lobule small to huge, may be uniform or variable in size on the same stem, sometimes covered by large stipules, semicircular in outline, inflated, rostrum obtuse, acute, hooked, or with a slender extension, straight and spinous, or curved, resembling an elephant's trunk. Cells ca. 20μ, basal larger, trigones small or confluent. Stipules very large to medium, usually broader than tall, margins may be uneven to bluntly toothed, apical sinus small, variously shaped, segments obtuse to acute. Cells may be different from those of the leaves. Invol. leaves broad, obtuse, the lower portion ventricose, completely enwrapping the perianth, upper portion undulate or spreading. Perianth subspherical, pedicelled, very smooth, sometimes with an apical depression, immersed or emergent. “The curious perianth of this species at first sight appears to be concrete with the involucral leaves, and looks far more like a naked calyptra than a perianth…”

Mitten's type is shown as having small lobules and the ventral base of the leaves broadly inflexed.

F. squarrosula” det. Stephani in Herb. Petrie, is F. patula.

North Island: On manuka, twigs, and bark in bush, near Atiamuri, tree trunks Puaiti Bush, manuka near Opepe, Mokoia near Hawera, bark in bush Brunswick Rd., Wanganui, K. W. A.; on manuka and tree trunks, 10 localities in Wairoa County, abundant on willows and cabbage trees, Otupae and Erewhon, 2–3,000 ft., bark in bush, Taihape, on bark, Esplanade Bush, Palmerston North, and Bledisloe Park, E. A. H.; trunks of trees not native. Mt. Bruce, Masterton, N. Welch; “Ngahere,” Puketitiri, E. A. H.; Mt. Climie. 2,000 ft. (Rimutakas), probably, (sterile), 1127, A. P. D.

South Island: Kennedy's Bush (Cashmere Hills), H. M. H.; near Mt. Cook, Hermitage, G. O. K. S.; Conical Hill (Otago), probably, V. D. Z., 17154, P. R. B. Herb.; on shrub, south of Dunedin, K. W. A.; on “pine” tree, Waihopai Scenic Reserve (Invercargill), 397, on Aristotelia, Bravo Is. (Stewart Is.), 411, W. M.: also Travers V. (Nelson), A. P. D.

Chatham Islands: R. Gilpin.

Form with spiniferous lobule is from the following places:

North Island: Beech forest, Rangitikei V. (Kaimanawas), A. P. D., 1032; tree trunk by L. Waikaremoana, on solitary manuka bush, Ohuka (Wairoa), beech bush, Tauruarau, on kanuka. Tauruarau Bridge (Napier-Taihape Rd.), E. A. H.

South Island: Cobb Valley (Nelson), A. L. H.; on beech tree, L. Roto-iti (Nelson), 2,100ft., G. O. K. S.; Castle Hill (Canterbury), H. M. H.; Happy Valley (Springfield), F. MacD.; on wineberry in forest remnant and tree trunk, Mt. Cargill (Dunedin), 398 in part and 402, W. M.; on manuka, Waitati (Dunedin), Field Club, per E. Campbell; Nugget Point, G. Simpson, 24705, P. R. B. Herb.

There are minor forms of this species which must be mentioned, with leaves rarely more than 0·5 mm. across; some may be at once recognisable as F. patula, others, of pale colour and with margins finely recurved, may look like a small F. subhampeana. The perianth when present remains constant and is that of F. patula, but often

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these little plants are sterile and gemmiferous with explanate lobules. Six of these small forms have been collected, within a ten-mile radius of Wairoa, also from bark by L. Waikaremoana on kanuka by Tauruarau Bridge, E. A.H.; in bush, Terapatiki (Waikaremoana), Mrs. G. O'Malley; on manuka, north-west L. Wairarapa Rd., H. M. H.; Kapiti Is., A. P. D.

The Handbook localities for F. patula are: Tarawera, on dead bark, Manawatu R., on fences, etc. Colenso.

Frullania kirkii St.

F. kirkii St., Hed., 145, 1894; Spec. Hep., iv, 407, 1910.

Plants robust, in dense mats or sprawling. Stems 4–5 cm. long, bi- or tri-pinnate, branches and branchlets usually short, basal ones often longer. Leaves imbricate, sub-circular, to 1·5 mm., concave, margins often narrowly incurved, dorsally broadly overlapping the stem, cordate at the base; lobules semicircular in outline, oblique, very large, to 0·7 mm. long on main stem, progressively smaller on the branches, inflated, dorsal side often wider than the ventral, rostrum almost always hidden, straight or hooked, acute or obtuse. Cells 18–20μ, trigones small. Stipules reniform to sub-circular, very broad on main stem, the largest on the type being 0·9 mm. wide, smaller ones to 0·5 mm., auricled at base, apex shortly bidentate, sinus small, often hard to detect. Invol. leaves very concave, appressed and almost covering the perianth, but exposing a narrow median slit, acute, lobule attenuated and may be twisted, lower margin more or less lacerate; stipules narrow with smallish lateral segments, main segments acuminate, decurved and involute. Perianth decurved, emergent, and thus appearing short and broad, no ventral keel, but may be tumid as the capsule develops.

Sterile plants of this species can scarcely be distinguished from sterile plants of F. patula Mitt.

North Island: On young beech, Whakapapa (Mt. Ruapehu), G. O. K. S.; on manuka twigs, Atiamuri, on larch in State Forest, 497, K. W. A.; on bark, Northern Ruahines, on exposed trunk of Fuchsia excorticata, hills west of Silverstream, (doubtful), trunks of manuka trees, river flat, Orongorongo V., H. M. H.; Akatarawa V., A. P. D., 925.

South Island: Twigs in bush, Maruia Springs (Nelson), J. H. McMahon; beech trees, L. Roto-iti, 2,100 ft., G. O. K. S.; trunks of beech trees, Arthur's Pass, W. M., 88 and 378; Avalanche Peak (Arthur's Pass), H. M. H.; on beech trees, Punch Bowl and Mingha V. (Arthur's Pass), Arthur's Pass, F. MacD.; Summit Road, Akaroa, (doubtful), H. M. H.; Travers V. (Nelson), 2,300 ft., A. P. D.; on rocks, summit of Saddle Hill, Dunedin, 380, W. M.

Stewart Is.: Log in bush, Oban, W. M., 393.

No. 239, Herb. Rodway, coll. Weymouth, labelled F. kirkii. det. Stephani, is F. deplanata.

The type is from the Bluff Hill, leg. Professor Kirk.

Frullania subdeplanata St.

F. subdeplanata St., Spec. Hep., iv, 408, 1910.

This is a difficult species. It appears to have the perianth and lobules of F. deplanata Mitt., and the broad round leaves and large, round to subreniform stipules of F. patula Mitt., and perhaps of

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F. kirkii St. Perhaps it is synonymous with F. scandens Mont, from Auckland Is., for a plant of this description has recently been collected on Campbell Is. by J. H. Sorenson. But one cannot be certain, as Montagne's plant was sterile. From the description of F. scandens in the Synopsis, either F. subdeplanata or F. kirkii or F. patula could be a synonym for F. scandens. Further specimens from the southern islands might throw further light on the matter.

Another fragment from Campbell Island has lobules resembling those of F. cranialis.

There are also border-line specimens between F. deplanata and F. subdeplanata and between F. subdeplanata and F. kirkii.

Specimens which are fairly satisfactorily accommodated under this heading are from the following places:

North Island: On manuka, E. of L. Taupo (perianths both with and without a ventral keel), K. W. A.; growing with a few stems of Glyphothecium alare, above Hopuruahine River, L. Waikaremoana, Gr. O. K. S.; on bark of small tree, Esplanade Bush, Palmerston North, E. A. H.; Mt. Egmont, with Lepidolaena sp., G. O. K. S.; on papery bark, watercourse, N. of Field Hut, Tararuas, 2,600 ft., V. D. Z., 7453, P. R. B. Herb.; Kapakapanui, 866, Wharite, 2,500 ft., A. P. D.

South Island: Arthur's Pass, F. MacD.

Frullania deplanata Mitt.

F. deplanata, Fl. Nov. Zel., ii, 161, 1855; Handb. N.Z. Fl., ii, 536, 1867; St., Spec. Hep., iv, 407, 1910; Rod., Tas. Bry., ii, 84, 1916.

Plants pale- or dingy-green, often rufescent, sometimes dark-redbrown, very common, widely creeping or compactly matted, on manuka, twigs, or bark of both small and big trees. Stems to 5 cm., but usually shorter, shortly alternately branched, in some forms shortly and densely branched. Leaves more or less glossy, contiguous to densely imbricate, oval to elliptic, obtuse, apices recurved sometimes to as much as ⅓ of the leaf; lobule with a semi-circular outline, upper swollen portion overlapping the stem, mouth about 0·15 mm. distant from the stem, and usually reaching below the ventral margin of the loaf. Cells usually rounded with confluent trigones; trigones sometimes minute, acute. In darker coloured specimens the cell walls are thick. Stipules very variable, sub-circular with uneven or even margins, to obovate or obcuneate with a shoulder angle or tooth; cells always irregular in shape with sinuous walls, thereby differing from the leaf-cells; apical sinus variable. Inmost invol. leaves ca. 1·3 mm. × 0·65 mm., acute to broadly acute, lobule linear with recurved margins, lower part often lacerate, stipule often lacerate, the leaves and stipule connate, to very much so. Perianth sometimes with a subfloral innovation, large obovate-oblong, convex, with lateral margins decurved, apex retuse, occasionally bent downwards as in F. kirkii, ♂ branches of varying lengths.

This species can generally be recognised by the smooth convex perianth minus a ventral keel, the lobules standing away from the stem, and the cells of the stipules being different from those of the leaves.

F. curnowii St. is a form usually pale green or faintly tinted, creeping on bark with longish stem, and leaves scarcely imbricate,

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F. grayana St., from vol. vi, Spec. Hep., is a compactly growing, much branched form, leaves somewhat larger.

Another compact form represented by a plant from bark, Ruakituri R. bed, also from Kapakapanui, 867, A. P. D., has the dorsal margin of the leaves finely recurved, 3 from Atiamuri, K. W. A., being somewhat similar.

From rock, edge of L. Waikaremoana, E. A. H., and from Terawhiti Hill, 820, A. P. D., are compact forms with stipules having recurved margins, but none of these forms, including F. grayana St., seems distinct enough to be a separate species.

North Island: Bay of Islands, V. W. Lindauer; Waitakeres (Auckland), E. D. Swanberg; National Park, G. O. K. S.; Ohakune Track (Mt. Ruapehu), 2, Waimarino V. (Kaimanawas), H. M. H.; Atiamuri, Taupo and Rotorua districts, mostly on manuka, 23 specimens, K. W. A.; Wairoa County, mostly on bark, including Waikaremoana and Morere, 17 specimens, Tauruarau and Taihape, 4, Bledisloe Park, Palmerston North, E. A. H.; Hopuruahine (Urewera), 2, Mt. Egmont, G. O. K. S.; Puketitiri (Hawke's Bay), M. Brownlie; Ruahine foothills, A. L. H.; Northern Ruahines, Orongorongo V., H. M. H.; Southern Ruahines, 752, Kapakapanui, 897, 886, beech forest, Rangitikei V. (Kaimanawas), 3,000 ft., 989, 1035, A. P. D.; Field Hut (Tararuas), V. D. Z.

South Island: Mt. Arthur (Nelson), L. Roto-iti, 2,100 ft., on beech bark, G. O. K. S.; Pelorus Bridge (Nelson), A. L. H.; Mt. Marina, V. D. Z.; Marlborough, 2, J. H. McMahon; Mingha V., Punch Bowl (Arthur's Pass), Castle Hill, 3, F. MacD.; Avalanche Peak, H. M. H., W. M., 377; Upper Waimakariri, M. Berry; near Cass Hut, Kennedy's Bush, Banks Peninsula, 4, Castle Hill, 3, H. M. H.; Castle Hill, in beech forest, L. B. M.; Waitati, Silver Peak, Mihiwaka, Glenledi, 24699, P. R. B. Herb., near Hindon (Central Otago), 2, G. Simpson; Maungatua Range, Berwick State Forest, K. W. A.; Mt. Cargill, Waihopai Scenic Reserve (Invercargill), 2, W. M.; Wakari, Field Club, per E. Campbell; on lacebark, Otago Pen., 406, W. M.

Stewart Island: On rimu, Pryce's Peak, 522, Little Glory Harbour, 427, W. M.

“East Coast and Interior, on bark, abundant, Colenso; Wellington, on lichens, Stephinson.”

Also recorded from Tasmania.

Sub-genus Diastaloba Spruce

Plants mostly small, often monoicous, lobules sub-cylindrical or pitcher-shaped, distant from the stem, mostly with a stylus intravening, and inclined away from it at an angle of not less than 30°, cells clear.

Sec. Graciles Verd.

Plants small, red-brown, leaves rounded or acute, mid-leaf cells not longer than 20μ.

F. rostrata (Tayl.), Syn. Hep.

Sec. Pictae Verd.

Plants small to medium, leaves with a basal vitta of cells often more discoloured than the rest, stipules toothed (in New Zealand).

F. aterrima (Tayl.), Syn. Hep.

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Sec. Lucidae Verd.

Plants robust, leaves taller than broad with a transverse insertion, cells of the mid-leaf not less than 20μ

F. ptychantha Mont.

Key to Species
1. Plants small to medium, mid-cells less than 20μ, lobules oblique 2
Plants robust, mid-cells greater than 20μ, lobules appearing pedicelled, angle of inclination rarely less than 45°, to 90° or, more ptychantha
2. Stylus conspicuous, stipules entire rostrata
Stylus reduced, stipules strongly laterally toothed aterrima

Frullania rostrata (Tayl.) Syn. Hep.

Jung. rostrata Tayl., Lond. Journ. of Bot., 87, 1845; Fl. Ant., 163, 1847. Frullania rostrata, Syn. Hep., 415, 1844–47; Handb. N.Z. Fl., ii, 538, 1867; Spec. Hep., iv, 659, 1910.

Frullania. dissitifolia St., Spec. Hep., iv, 659, 1910.

Plants small, usually dioicous, sometimes monoicous, reddishbrown, sometimes darker or paler, occasionally greenish or green, corticolous, rarely on rocks, sometimes climbing through tufted bryophytes. Stems to ca. 3 cm., usually shorter, subpinnately or, in elongated forms, regularly pinnately branched, pinnae ca. 2 mm., sometimes longer with 1–2 pinnules. Stem leaves imbricate to remote, the latter may be 0·6 or 0·7 mm., but usually 0·4, on very slender stems even smaller, occasionally caducous, oval or oblong-oval, concave, apices incurved, sub-acute to apiculate; lobules oblong-pitcher-shaped, oblique, tilted somewhat with the mouth directed inwards, mouth often hyaline; stylus intravening between it and the stem, triangular to ca. 0·1 mm. tall, or reduced to a stylate tooth. Branch leaves similar but smaller, more crowded, more acute; lobules beautifully regular, not varying in size from those of the stem. Cells rounded-oblong, mostly all separate and usually in rows, becoming shorter and smaller towards the apex, and larger at the base, but not forming a distinct vitta as in F. aterrima. Stipules oval, may be broadly oval or circular, entire, or perhaps with an: undulation, or scarcely toothed, deeply bi-fid, segments obtuse to sub-acute, erect or slightly converging. Invol leaves oblong, incurved, apiculate, margins of lobules and stipules reflexed, entire. Perianth obovoid to narrow obovoid, normally triquetrous, dark to blackish, shining, faces often convex, but may be plicate with minor ridges. ± bracts in pairs from 2·6, on special, short ♀ branches.

F. congesta (Tayl.), Syn. Hep., the description of which appeared a year earlier, as represented by the type from Auckland Is., is a difficult species to deal with. It is closely related to F. rostrata, in that the stipules, lobules and colour are the same. The cells, however, are bigger than is usual in F. rostrata, and resemble in appearance those of F. ptychantha. The leaves, many of which have fallen off, are definitely imbricate and convex, but vary greatly in size, and stems and branches may bear both small and large (comparatively) ones. A plant from Stewart Island (W. M., 362) might be placed here, as it has leaves all equally imbricate, and similar cells.

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The Handbook and Fl. Novae Zelandiae both record it from the mainland, but specimens nearest to it in appearance almost invariably turn out to be F. aterrima.

Localities for F. rostrata are as follows:

North Island: On tree trunk in manuka, on bark with Mastigophora, on manuka on hillside, Waipoua Forest, K. W. A.; on kauri bark, Waipoua Forest, Rangitoto Is., upper branches of totara, Pukemako (King Country), L. B. M.; Little Barrier Is., on Lepicolea sp., W. M. Hamilton; on taraire bark, Coromandel Range, A. L. H.; Ohakune Track, 3,000–4,000 ft., H. M. H.; on bark, near Ohakune Hut, 4,500 ft., mostly on bark, Whakapapa, 4 specimens, G. O. K. S.; amongst mosses on trees near Chateau, National Park, 3,000 ft., on manuka bark, near Atiamuri, 18 specimens, on manoao, near Opepe, on larch, Rotorua State Forest, on rotten log, Mangarewa Gorge (between Rotorua and Te Puke), on manuka, E. of Waiotapu and Kaingaroa Plains, on pine tree, Kaingaroa, on trunks of red beech, Pukerimu Bush, small tree in bushy gully, between L. Roto-iti and Coast, K. W. A.; Raukokore R. (B. of Plenty), G. O'Malley and G. O. K. S.; Maungapohatu (Urewera), B. Teague; Summit Ridge, Mt. Manuoha, G. O. K. S.; climbing on Chandonanthus squarrosus, on log, Waikare-iti, 3,000 ft.; plentiful on bark round L. Waikaremoana, Morere Bush, on bark, 2 specimens, bush at Maungapoike Falls (Wairoa), one on upper branches of tawa tree, 3 specimens, E. A. H.; on rocks of crater rim, amongst mosses, Mt. Tongariro, Mt. Egmont, L. B. M.; on bark in beech forest, Rangitikei V. (Kaimanawas), 3,000 ft., 13 specimens, A. P. D.; Birch Range (Kaweka), ca. 3,000 ft., E. S. West; beech bush, Tauruarau (Napier-Taihape Rd.), plentiful, on bark in bush, Taihape, E. A. H.; Wharite, 2,000 ft., A. P. D.; Peninsula (Tinui), R. W. Hewitt; Tauherenikau, Wairongomai R., 7236, V. D. Z.; Kapakapanui, 875, 861, 863, Akatarawa V., 926, Upper Hutt, 915, Mungaroa, 685, A. P. D.; on manuka, Southern Rimutakas, over 3,000 ft., on manuka, north-west L. Wairarapa Rd., on manuka trees on river flat, Orongorongo V., H. M. H.; Wilton's Bush, R. Mason.

South Island: Marlborough, 2 specimens, J. H. McMahon; on beech bark, L. Roto-iti, 2,100 ft., G. O. K. S.; Pelorus Bridge (Nelson), A. L. H.; Arthur's Pass, 4 (Bealey Track, Avalanche Peak), H. M. H.; Mingha V. (near Arthur's Pass), F. McD.; beech trunk, near Avalanche Crk. (Arthur's Pass), 95, W. M.; Mt. Cook, G. O. K. S.; near Fox Glacier, Mrs. Knight; rock at Leith Stream (Otago), Leith Saddle near Dunedin, W. side of Mt. Maungatua, on bark of Weinmannia sp., Glenledi, G. Simpson and J. S. Thomson; tree trunk in bush, between Taieri and Akatore Rs., on rocks, Maungatua, ca. 2,500 ft., K. W. A.; Mt. Maungatua, 24693, P. R. B. Herb (in part), west side of Mt. Maungatua, Silver Peak, G. Simpson.

Stewart Island: Bark of rimu, Pryse's Peak, 521, on dead log, Glory Harbour (Paterson's Inlet), 395, on deciduous scales of miro, Garden Mound, 342, W. M.

Auckland Island: On Radula physoloba (minute), 2279, Prof. Du Riete, comm. Dr. H. Persson. Another small specimen from Campbell Island, coll. J. H. Sorensen, resembles this in appearance, but has large basal and mid-leaf cells and a rugulose perianth.

The type was from Auckland Is., presumably collected by Hooker,

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Frullania aterrima (Tayl.), Syn. Hep.

Jung. aterrima Tayl., Lond. Journ. of Bot., 395, 1844; Fl. Ant., 1855.

Frullania aterrima, Sym. Hep., 450, 1844–47; Fl. Nov. Zel., ii, 162; Handb. N. Z. Fl., ii, 538, 1867; Spec. Hep., iv, 657, 1910.

Plants monoicous or dioicous, usually compact, olive-green to dark-purplish-brown, rarely red-brown. Stems usually to ca. 2 cm., branching pinnate, bi-pinnate or irregular, dense or lax, branches very variable in length, sometimes increasing in width upwards and may be slightly curved. Leaves generally bigger on the main stem, and less imbricate, concave, oval to broadly oval, rounded at apex, or occasionally acute, rarely caducous; lobules similar to those of F. rostrata, but may be more oblique on the main stem, constant in size, usually hyaline at or near the mouth; cells ca. 18μ, minute towards the upper margin, suddenly enlarged at ventral base to form a vitta with 6–sided cells, to ca. 35μ, which is usually visible with the leaves in situ, but may be somewhat hidden by the broad stipules. Stipules 3–4 times as broad as the stem, with at least 1 prominent lateral tooth, deeply bi-fid, segments widely diverging. Invol. leaves and perianth as in F. rostrata, also ♀ inflorescence.

May be distinguished from F. rostrata by the broad, laterally toothed stipules with gaping sinuses, also by the cluster of large cells at the ventral base, and the usually darker colour.

North Island: Low epiphyte, kauri bark, Waipoua Forest, L. B. M.; trunk of living kauri, 6 ft. from ground, Waipoua Forest, on manuka and manoao, near Atiamuri, including nos. H147, H91, H197, 8 specimens, on bark in bush, Rotorua-Atiamuri Rd., on manuka, E. of Taupo, on manuka, E. of Waiotapu V., K. W. A.; on bark, Whakapapa, 2, bark of young beech, Whakapapa, with F. rostrata, National Park, near Tangiwai (Main Trunk), Panikau, G. O. K. S.; Ohakune Track (Mt. Ruapehu), Waiotaka V. (Kaimanawas), H. M. H.; with F. rostrata, Mt. Maungapohatu, B. Teague; on bark, edge of L. Waikaremoana, 2,000 ft., Tauruarau, on beech bark, E. A. H.; Northern Ruahines, Oroua (Southern Ruahines), Southern Rimutakas, on bark of leatherwood bushes, Tararuas, 3,500 ft., 960, manuka trees on river flat, Orongorongo V., H. M. H.; Wharite, 2,500 ft. (Southern Ruahines), 3, Southern Ruahines 765, 745, Kapakapanui, 854, 865, on rocks with Herberta alpina Mt. Holdsworth (Tararuas), 4,100 ft., 947, Akatarawa V., 929, A. P. D.; Mt. Climie, 2,000 ft., Rimutakas, 2 gatherings, A. P. D.

South Island: Travers V. (Nelson), 2,300 ft., A. P. D.; Punch Bowl (Arthur's Pass), F. MacD.; Arthur's Pass, H. M. H.; Avalanche Peak Track, Arthur's Pass, H. M. H. and F. MacD.; on Aristotelia fruticosa, 2 miles E. of Arthur's Pass township, 130, on tree-trunk, Kelly's Hill (Otira), with F. incumbens, 379, on wineberry, Mt. Cargill (Dunedin), W. M.; on tree-trunk in bush between Taieri and Akatore Rs., ca. 600 ft., on tree-trunk in bush, saddle between Waitati and Dunedin, ca. 1,000 ft., K. W. A.

The type was from Auckland Is., coll. Hooker.

All the variations of F. rostrata appear to have their counterparts in F. aterrima, though the elongated form with contiguous to remote leaves is rare. (One such specimen from Rangitikei V., 3,000 ft., 968, A. P. D., is green.)

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There is a neat, tiny form, here described as var. lepida var. nov., which can hardly be passed by, as it has cropped up in several places, with perianths distinctive-looking with a median dorsal furrow.

Var. lepida var. nov.

Minima, pallida olivacea, caulis ramosus, 0·5 cm. longa. Folia caulinis ramorumque aequiparva, ca. 0·2–0·3 mm., imbricata, involucralia appressa, perianthia oblonga-obovata antica facie unisulcata, postica late carinata.

High land, E. of Waiotapu V., on manuka, Kaingaroa Plains (near Murupara), ca. 1,500 ft., H501, 24/6/34, trunk of small tree in bush gully between L. Rotoehu and coast, 12/5/43, K. W. A.; National Park, G. O. K. S., 1932; with green F. rostrata, beech forest, Rangitikei V. (Kaimanawas), 969, Jan., 1947, Kapakapanui, Nov. 24, 1946, 860, A. P. D.; Wilton's Bush (Wellington), Oct., 1941, R. Mason; L. Roto-iti, South Island, 2,100 ft., G. O. K. S.

The specimen from Kapakapanui is dark-red-brown.

Though the type of F. aterrima must be considered a small form of a variable species, this little variety is noticeably different.

Another striking form of F. aterrima has the appearance (numerous small branches), and sub-horizontal to horizontal stem lobules (branch lobules more normal), of F. ptychantha, but the toothed stipules and leaf-cells, including the vitta, are definitely those of F. aterrima. This specimen, off tree stems at Te Wharau, near Carterton, Sept., 1942, no. 407, W. M., could well be the result of a cross between F. aterrima and F. ptychantha.

Frullania ptychantha Mont.

F. ptychantha, Mont., Annal. des Sc. Nat., 19, p. 258; Voy. au Pole Sud, 1, 225; Syn. Hep., 442, 1844; Fl. Nov. Zel., ii, 163, 1855; Handb. N.Z. Fl., ii, 577, 1867; St., Spec. Hep., iv, 662, 1910. Jung, ptychantha Hook., Crypt. Ant., p. 51. Jung, myosota Tayl., Lond. Journ. of Bot., 393, 1844.

Plants olive-green to light-red-brown. Stems to 4 cm., all lying parallel, closely pinnately branched, branches short to very short, often bi-pinnate. Leaves sub-rotund from a somewhat narrowed base, to 1·3 mm., sometimes a little broader than tall, much smaller on branches, exappendiculate with a transverse insertion; cells ca. 30μ, 6-sided with acute trigones, or rounded with trigones all confluent, basal to 45μ lobules small, uniform in size, to about 0·15 mm. long, sub-symmetrical, narrowed to the mouth, distant from the stem, appearing pedicelled, very oblique, occasionally reaching below the horizontal. Stipules sub-rotund or more usually obovate, sometimes shouldered, bi-fid to almost ½, lobules diverging. Inmost invol. leaves acute or apiculate, a little falcate, entire or with an occasional tooth, 0·5 mm. broad, “lobules linear-lanceolate, entire acuminate” (K. W. A.). Stipule deeply bi-fid, segments narrow-triangular, attenuate. ♂ branches very short with only 1–2 pairs of bracts (in plants seen). Perianth variable, commonly with 1 dorsal and 2 ventral keels, but supplementary keels may bring the number up to 9, as in the earlier descriptions.

Distinct from the other New Zealand species in the small, distant, oblique lobules, the very short closely pinnate branches, and probably in the exappendiculate leaves.

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North Island: Epiphytic in tawa forest, ca. 1,500 ft., Moehau, Summit Rocks, Moehau, H22, amongst mosses on ground, Summit, Te Moehau, epiphytic and firmly attached to Podocarpus totara in exposed position, ca. 2,000ft., Mt. Moehau, L. B. M.; Mt. Archeria (Little Barrier Is.), ca. 2,300 ft., B. E. N. Matthews; scrambling amongst other hepatics, South Rimutakas, H. M. H.; on bark, Kapakapanui, 849, with Goebeliella cornigera, South Ruahines, 739, A. P. D.; Waiopehu Ridge (Tararuas), G. O. K. S.; Mt. Climie, 2,000ft. (Rimutakas), A. P. D.

South Island: Lead Hills near Bainham (Nelson), G. Simpson and J. S. Thomson; forest trees, Bluff Hill, 450, on trunk of fallen tree (near top), Garden Mound, Stewart Is., 341, the commonest species on trees, Table Hill forest, Stewart Island, 498, on “kamahi” bark, Paterson's Inlet, Stewart Is.; 376, W. M.

Campbell Is.: J. H. Sorensen.

“North and South Is., Wellington, Stephenson, Lyall; Lord, Auckland and Campbell Is., Hombron and Hooker.”

In addition to the types, a full set of New Zealand species of Frullania has been deposited in the Herbarium of the Botany Division of the Plant Research Bureau. It is with much pleasure that F. allanii has been named in honour of Dr. H. H. Allan, the head of this Division.

Excluded Species.

F. hypoleuca Nees, Handbook, p. 537, is a Hawaiian and Indo-Malayan species. The plants referred to this species by Hooker are most probably F. ptychantha. F. hypoleuca has a papillate lobule.

F. magellanica (Spreng.) Web., Handbook, p. 538, is a South American plant with minutely denticulate leaves. Apparently not a New Zealand species.

F. gracilis Nees, Handbook, p. 539, a widely distributed species, but not represented in New Zealand. The involucral leaves are irregularly toothed and the stipules quadrate.

F. cornigera Mitt., Handbook, p. 535, now Goebeliella cornigera (Mitt.) St., monotypic, and in a family of its own, is a very notable and much-prized species. It is a robust, olive-brown plant with much the appearance of a Porella from the dorsal aspect. Leaves with hyaline margins and a double lobule, stipules quite entire. It has recently been collected in both Little Barrier and Auckland Is., otherwise peculiar to New Zealand.

Colenso's Species.

In addition to those species of Colenso's already mentioned, Stephani reduced 3 more as follows:—

F. minutissima Col., Trans. N.Z. Inst., 19, 1886, p. 298

F. delicatula Col., Trans. N.Z. Inst., 19, 1886, p. 297}

= F. falciloba juvenile form

F. minutissima Col. (another specimen) = F. rostrata

Others not mentioned by Stephani are:

F. scabriseta Col., Trans. N.Z. Inst., 19, 1886, p. 298

F. amplexicaulis Col., Trans. N.Z. Inst., 18, 1886, p. 299

F. tongariroense Col., Trans. N.Z. Inst., 21, 1888, p. 73

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F. intermixta Col., Trans. N.Z. Inst., 21, 1888, p. 74

F. platyphylla Col., Trans. N.Z. Inst., 21, 1888, p. 74

F. curvirostris Col., Trans. N.Z. Inst., 21, 1888, p. 76

F. polyclada Col., Trans. N.Z. Inst., 21, 1888, p. 76

Without the types it is difficult to say with certainty what these species really are.

Additional Localities. *

New and interesting localities of species recently received are as follows, the subantarctic specimens being mostly from coastal scrub and forest, collected by Professor G. Einar Du Rietz and Mr. H. Guthrie-Smith in the autumn of 1927.

F. pentapleura: Log on ground, bush remnant, North Otago.

K. W. A. 930, 20/7/48 (gemmiferous).

F. squarrosula: Rangitoto Is., L. H. Millener; on oak-tree, Wallaceville, A. P. D. 1134.

L. falciloba: On ground in the tussock country, Antipodes Is., Du Rietz 2597.

F. rostellata: Onetangi Beach bush, Waiheke Is., A. L. H. 1345.

F. deplanata: Auckland Is., Guthrie-Smith 2628 comm. H. Persson; Campbell Is., Du Rietz 2578.

F. subdeplanata: Auckland Is., Guthrie-Smith 2628 comm. H. Persson; 2279 (± branches with over 20 pairs of bracts), 2239, 2297; Campbell Is., 2566, Antipodes Is., 2597, Du Rietz.

F. ptychantha: Auckland Is., 2238 Du Rietz.

F. rostrata: Campbell Is., 2580, 2576, 2578 Du Rietz; Auckland Is., Guthrie-Smith 2322 comm. H. Persson.

F. congesta: Auckland Is., 2243, 2292, Du Rietz.

F. aterrima: Auckland Is., 2298, 2301, 2293, Du Rietz; 2322 comm. H. Persson, Guthrie-Smith.

It seems now almost certain that F. subdeplanata St. is the same species as Montagne's F. scandens.


The writer wishes specially to thank the collectors who have so abundantly provided the material about which this paper is written. Grateful thanks are also tendered to Sir Edward Salisbury, of Kew, and Professor Charles Baehni, of Geneva, for the loan of Taylor's and Stephani's types respectively; to Miss Curtis, of the Royal Society of Tasmania, for the loan of Rodway's specimens; to the authorities of Sydney and Melbourne Botanic Gardens for the use of literature and specimens; and to Mr. A. D. Banwell, Bibliographer of the British Bryological Society, for help and advice in the procuring of literature. The gift to New Zealand of fragments of Mitten's types by the late Dr. Marshall Howe, of New York, and the much-valued assistance rendered in the early stages by the late W. E. Nicholson, of the British Bryological Society, have already been acknowledged in earlier papers.

A further acknowledgment is of the loan of Pearson's syntypes from the Herbarium, Manchester University, kindly sent by Mr. J. G. Roger.

[Footnote] * Also F. cranialis from “Otupae,” N.W. Ruahines, A. P. D., Dec., 1947, which is the first recording of this species from the North Island.

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Bastow, R. A., 1887–88. Tasmanian Hepaticae, Papers and Proc. Roy. Soc. Tasmania, pp. 209 289, 1887, pl. 1–34, 1888. Hobart.

Colenso, Rev. W., 1880. A Description of Some Newly Discovered Cryptogamic Plants. Trans. N.Z. Inst., vol. 19, pp. 200–299.

—— 1887. Trans. N.Z. Inst., vol 20, p. 252.

—— 1888. Trans. N.Z. Inst., vol. 21, pp. 73–77.

Cooke, M. C., 1907. Handbook of British Hepatics. Edinburgh.

Gottsche, C. M., Lindenberg, J. B. G., et Nees, C. G., ab Esenbeck, 1844–47. Synopsis Hepaticarum, including Supplement. Hamburg.

Hooker, Sir J. D. The Botany of the Antarctic Voyage, 1839–1843. London.

—— cum, W. Mitten, 1855. The Botany… Flora Novae Zelandiae, vol. 2.

—— cum T. Taylor, 1847. The Botany… Flora Antarctica, vol. 2.

—— cum W. Mitten, 1860. The Botany… Flora Tasmaniae.

—— cum. W. Mitten, 1867. Handbook of New Zealand Flora, vol. 2. London.

Nicholson, W. E., 1925. Notes on Some New Zealand Species of Frullania. The Bryol., vol. 28.

Pearson, W. H., 1923. Notes on a Collection of New Zealand Hepatics. Univ. Cal. Pub. Bot., vol. 10, p. 317.

—— 1924. Notes on Tasmanian Hepatics. Kew Bull., Misc. Inf., no. 2, pp. 68–75.

Rodway, L., 1916. Tasmanian Bryophyta, Hepatics. Papers and Proc. Roy. Soc. Tasmania.

Sim, T. R., 1926. The Bryophyta of South Africa. Trans. Roy. Soc. of South Africa. Capetown.

Spruce, R., 1849–62. Hepaticae Amazonicae et Andinae. Trans. Bot. Soc. Edinburgh.

Stephani, Fr, 1892. Colenso's New Zealand Hepaticae. Journ. Linn. Soc., vol. 29, pp. 263–280. London.

—— 1911. Species Hepaticarum, vol. 4. Geneva.

—— 1924. Species Hepaticarum, vol. 6, Geneva.

—— Hepaticae Species Novae, 1865–96. (Reprints from issues of Hedwigia.)

—— Icones Hepaticarum. Unpublished Hand-drawings of Species.

Verdoorn, Fr., 1929. De Frullaniaceis IV and V. Annales Bryologici, vol. 2, pp. 117–154. vol. 2, pp. 155–164. The Hague.

—— 1930. Die Frullaniaceae der Indomalesischen Inseln. De Frullaniaceis VII. Ann. Bry., 1930, Supp. Vol. The Hague.