New Zealand Hepaticae (Liverworts)—XI
A Review of the New Zealand Species of the Genus Lepidolaena
[Received by the Editor, December 18, 1958.]
Description. of genus with 3 new sections, Lepidolaena, Dolicocarpa, Dolicotricha—key to species—8 species described with 1 new species L. Berggrenii—L. brachyclada excluded from New Zealand.
I am indebted to Dr. R. E. Hatcher, of Illinois, for the information that Trevisan (1877) segregated L. Magellanica into a new genus Gackstromia on the basis of its peculiar fruiting structure. As already described, this structure is glabrous with a piliferous mouth, with ± sheathing floral leaves, also piliferous. I think it is no more generically distinct than is L. Menziesii, with paraphyllous stems and a long, rigid tubular coelocaule with the wall 8 cells thick. In other respects both of these taxa are essentially lepidolaenoid. After much thought, I prefer to keep the original genus intact with 3 sections. As Article 60 of the Code does not make it mandatory for authors to retain the existing name when changing the rank of the taxon, I retain the section as sec. Dolicotricha in preference to the ugly sounding name of Gackstromia.
Lepidolaena is a genus of the Southern Hemisphere excluding South Africa, consisting of 14 species, 8 of which are found in New Zealand and the subantarctic islands, 4 of these being described by Hooker senr. (1818–1820) in his Musci Exotici, 3 as new species under the generic name of Jungermannia. In 1835 Dumortier briefly described the unusual inflorescence under the heading of Lepidolaena. He then listed 3 of Hooker's species, L. claviger, L. palpebrifolia and L. Menziesii without any comment and with no reference which would identify them. L. Magellanica, the earliest of the species to be discovered and described, was apparently unknown to Dumortier. In spite of the inadequate description and the invalid combinations, the name of Lepidolaena is considered valid and remains inviolable. Lindberg (1873) drew attention to Dumortier's little known generic names and the fact that they must take precedence over later names then in use.
The position of Lepidolaena within the present framework of taxonomy is obscure. The old family Ptilidiaceae to which it belonged with 14 other genera, including Trichocolea, is in process of being dismantled. A new suborder, Ptilidiinae, has been inaugurated (Schuster, 1958), and new groupings in new families are emerging, but it is doubtful if a satisfactory arrangement will ever be arrived at.
Trichocolea has already been segregated by K. Muller (1945, p. 586) followed by R. Hatcher (1958, p. 237), who states that Trichocoleaceae is based upon the development of a coelocaule or stem perianth, and the absence of a true perianth. If this is an adequate basis for the segregation of this family, then Lepiodolaena must also be included in Trichocoleaceae, as the species of this genus have stem perianths or coelocaules, though in different forms. Also the leaves of L. Menziesii with their long, articulated cilia strongly resemble the leaves of Neotrichocolea and Tricho-
coleopsis, 2 genera from Japan. Also, I learn from Dr. S. Hattori of that country, that the long coelocaule of L. Menziesii resembles that of Trichocolea tomentella and Neotrichocolea Bissetti, while the stem perianths of other species resembles those of Trichocoleopsis sacculata, which has saccate lobules to the leaf, though these are not clavate as in Lepidolaena and the subgenus Diastoloba of Frullania. On the other hand, Lepidolaena has incubous leaves, while Trichocolea has them succubous.
The forms of the coelocaule in Lepidolaena are so diverse that I have felt obliged to divide the genus at least into sections, though the idea of new genera has not been discountenanced. These 3 new sections are as follows:
Lepidolaena sec. nov.
Planta parva vel magna, caulis nudus, coelocaule non elongato, dense paraphyllifero.
L. clavigera, L. palpebrifolia, L. Taylori, L. reticulata, L. Berggrenii, L. allophylla probably, as inflorescences are unknown.
Dolichocarpa sec. nov.
Planta robusta, pallida, interdum brunnescentia, caulis paraphylliferus, coelocaule elongato, ad 8 mm longo, infra tubulato, pauciparaphyllifero, superne latiore.
Dolicotricha sec. nov.
Planta robusta, rufo-brunnea, caulis nudus, coelocaule non elongato, nudo, folia floralia plurijuga, concava, dense longeque pilifera, pseudoperianthium formantia.
Lapidolaena Dumortier Recueil d'Observations sur les Jungermanniacées Fasc. 1, 1835.
Polyotus Gottsche in G. L. et N. Syn. Hep., 244, 1845.
Plants dioicous (in New Zealand), minute to robust, usually sterile, variously coloured, usually scrambling on or through other vegetation or creeping on bark. Stems short to very long, much branched, branches from the axils of the leaves, regularly short or with long ones intervening, or as subfloral innovations of unlimited length; cortical cells oblong to quadrate. Leaves incubous, bilobed, of the stem, remote to slightly imbricate, plane or amplexicaul, entire or toothed, ventral lobe or lobule usually saccate, often conjoined with a spinous lamina; leaves of the branches smaller, often more toothed or ciliate than the cauline. Cells small with thick walls (in L. Magellanica) to fairly large, trigones various, cuticle smooth or lowly papillose. Underleaves bi- or quadrifid on the main stem with margins of the segments entire or toothed; on the branches the segments are both plane and saccate, often reduced to 2 only, rhizoids when present arising from a basal lamina. ♀ Inflorescence terminal on a primary stem, in the fork of 2 branches rising at an acute angle; floral leaves and underleaves dense, large and irregularly shaped; archegonia numerous at the extremity of the stem, infertile ones carried upward with the growth of the calyptra which is adnate with a prolongation of the stem tissue, surrounding the developing sporophyte, this perigynium forming a coelocaule, of 3 different types (in New Zealand), which is naked or vested sparingly or densely with entire or toothed paraphylls. ♂ inflorescence in pairs along a ♂ branch, or terminal in 2 or 3 pairs on vegetative branches; bracts bilobed, conduplicate, lobules inflated, subequal to the lobes, antheridia single, not always present.
A distinct genus of the Southern Hemisphere (except South Africa), containing 14 described species, 8 of which are in New Zealand and the subantarctic islands, and 2 more in Tasmania (one undescribed).
Key to Species
|1.||Stem underleaves 4-fid, deeply divided, segments with margins flat or recurved, toothed, ciliate or entire||2|
|Stem underleaves shortly 2-fid, broad as in Frullania||7|
|2.||Plants pale to dingy, stems paraphyllous, densely bipinnate, with pinnae shortening in length towards the apex of the stem, leaves and underleaves everywhere closely and longly ciliate, coelocaule with a very long neck||Menziesii|
|Plants not pale (except L. claviger which is often pale glaucous green) dull green, greenish brown to reddish brown, or|
|crimson purple to blackish, stems without paraphylls, coelocaule pyriform to obovate||3|
|3||Stems with short branches, longer ones present in L. claviger, underleaf segments with entire margins, coelocaule paraphylls mostly entire||4|
|Robust, stems with both long and short branches, underleaf segments with spinous-toothed margins, coelocaule paraphylls with toothed margins||6|
|4||Leaves usually pale glaucous, of firm texture, occasionally with tints of rose or purple, ventral leaf-bases entire, margins of underleaf segments recurved, when 2 segments are present, they are median, not lateral, margins of leaf-lobules showing as crenulato-papillose||clavigera|
|Plants and leaves small, surface of lobules smooth or nearly so, segments of underleaves flat, obtuse or tapering, if 2 saccate segments are present, they are lateral and not median, pinnae of stems short, underleaves of the pinnae may be reduced to 2 segments, leaf cells large||5|
|5||Leaves glossy, leaves of pinnules narrow, almost always toothed, creeping on bark||Berggrenii|
|Leaves not glossy (in specimens seen) leaves of branches and branchlets (pinnae and pinnulae) not toothed, or if so only a little at the apex||reticulata|
|6||Leaves, specially the stem leaves, with toothed to laciniate-toothed bases, but otherwise mostly entire||Taylori|
|Branch leaves all ciliate-toothed all round, stem leaves partly so, the upper portion usually entire, texture soft||palpebrifolia|
|7.||Plants robust, reddish brown, leaf margins spinous-ciliate, stem underleaves with reflexed margin, a little toothed laterally and at the base at times, coelocaule smooth, regularly enveloped with floral leaves||Magellanica|
|Plants minute, dark red or purplish, stem leaves not ciliate, but spinous teeth present on the branch leaves, mainly round the apex||allophylla|
Lepidolaena clavigera (Hook senr.) Trev. ex Dumort.
Jungermannia clavigera Hook. senr. Musci Exotici I tab. 70, 1818.
Lepidolaena clavigera Dumort. Rec. d'Obs. sur les. Jung. p. 13, 1835, nom nud.; Trev. Sch. di una class. delle epatiche, 393, 1877; Steph Spec. Hep., 46, 1909.
Polyotus claviger G. L. et N., Syn. Hep., 245, 1845; Mitt., in Fl. Nov. Zel. ii, 152, 1854; Hook. Handb. N.Z. Fl., ii, 529, 1867.
Lepidolaena Stangeri Hodg. in Martin Bry. of Stewart Is., Trans. Roy. Soc. N.Z., 1950.
Plants dioicous, variable, compact or straggly, on bark or twiggy shrubs, variously coloured, mostly glaucous drying out to whitish. Stems to 6 cm, but often shorter, tufts of brown rhizoids often present at the base of the underleaves, regularly to irregularly bipinnately branched, pinnae short, usually 2 mm, but may be longer, close or distant, regular or with longer ones at intervals, the short pinnae horizontal. Leaves of firm texture, of the stem, imbricate, to 1.05 mm, cordate-ovate, appressed or partly so, lifting from the stem along the dorsal (upper) margin, acute to apiculate, entire or maybe a little toothed at the apex; lobules variable, explanate or clavate-saccate, joined and adjacent to a variously shaped, sometimes cucullate lamina, papillose with convex cell-walls, inclined from the stem from the base upwards, with or without a lateral spine; of the branches, smaller, generally toothed, lobules clavate-saccate, with or without a spine, inclined from the stem, often appearing pedicelled by reason of the reflexed ventral margin of the leaf. Underleaves, of the stem, 4-fid to about ⅔ of the length, segments lanceolate, acute, margins entire, rarely a little toothed, reflexed, occasionally the 2 median ones may be saccate-clavate; of the pinnae (branches) 4-fid, the 2 median ones saccate-clavate. Cell cavities 20–25μ regularly oblique, often indistinct, basal larger. ♀ Inflorescence terminal, usually with branches springing from a little lower down on the stem, pyriform, 2–4 mm; floral leaves toothed like the branch leaves, some longer, lobule mostly elongate toothed; underleaves 4-fid, margins reflexed and laciniate-toothed; coelocaule sparsely or densely covered with paraphylls; flimsy, 2 or 3 cells thick, but strengthened by the numerous paraphylls, cells vaguely defined, sterile pistillidia at the mouth; paraphylls short to elongate, concave, short and long ones interspersed, straight or decurved, mostly entire, cells large to 40μ clear, marginal ones smaller; capsule valves 1.5 mm long by 0.7 mm broad, 2 cells thick, margins slightly winged with marginal cells showing as transversely striate; seta short, cross section roughly hexagonal, 0.3 mm, white, opaque; elaters ca. 0.2 mm
long, closely bispiralled. In one instance 3 mature inflorescences were found at the extremity of one stem. Androecia on shortish branches, straight or decurved, bracts in 5–6 pairs, closely imbricate, subequally bilobed, keeled as in Lejeunea or Radula, lobule inflated, apex acute, antheria apparently sessile, to 0.15 mm in diameter.
In a species so comparatively common and well distributed as L. clavigera, it is only to be expected that there will be some aberrations from the type. For instance, one specimen from Stewart Island has the cauline leaves on some stems toothed all round. In another, all the segments of the cauline underleaves are clavate-saccate, and all the lobules, giving the ventral side of the plant a most singular appearance. In elongated stems the leaves may be scarcely imbricate, and the branches distant and irregular. But in the main the species is easily recognized by the characteristic firm texture of the glauco-whitish leaves, the short horizontal pinnae, the cauline underleaves with margins of the segments reflexed and entire, the entire paraphylls on the inflorescences when this is present, and the crenulate margins of the lobules.
Var. Stangeri Nees of the Synopsis Hepaticarum has the branch leaves toothed, and this is common, but most specimens have at least some of the branch leaves at least partially dentate.
North Island: Whangarei Heads, 24714 L. B. M.; Otaua Papakauri 4221a, Ohaeawai, Tauranga, 4228 S. Berggren; Waipoua Forest H781, H783; near Atiamuri, H205; Mt. Ngongataha, 5855; Mt. Messenger, 5822, K. W. A; Rangitoto Island, 5845, L. H. Millener; Great Barrier Island, 5845, R. Medland; Ohura 10222, 10201; Waikaremoana, 5807; Makaretu Stream, 5821; Silverstream Bush, 9765, E.A.H.; Kapiti Island, 5796; Mt. Kapakapanui (Tararuas), 898, A.P.D.; Stratford Park, 9703, W. M.; South Rimutakas, H. M. H.
South Island: Marlborough, 82, 10379, J. H. McMahon; S. of Hokitika, 1574, G. Einar Du Rietz; Otira, 4223, 4225, S. Berggren; Paparoa Range, H. W. Wellman; Fox Glacier, 5820, Mrs. Knight; Milford Track, 1623, F. Macdonald; Bligh Sound H868; Doubtful Sound, G. S.; Franklin Mts., W. A. Thomson; Waiho, 9811, L. Manapouri, 5632; Longwood Range, 6 specimens; Mt. Cargill, 403; Otago Peninsula, 406; Stewart Island, 12 specimens, W. M.
The type was from Dusky Sound, coll. Menzies. Endemic as far as is known.
Lepidolaena palpebrifolia (Hook. senr.) Trev. ex. Dumort.
Jungermannia palpebrifolia Hook. senr. Musci Exotici i tab. 71, 1818.
Lepidolaena palpebrifolia Dumort. Rec. d'Orbs. sur les Jung., p. 13, 1835 nom. nud Steph., Spec. Hep., iv, 49, 1909.
Polyotus palpebrifolius G. in G.L. et N., Syn. Hep., 246, 1845, Mitt. in Fl. Nov. Zel., ii, 246, 1854; Hook. Handb. N.Z. Fl. ii, 529, 1867.
Plants palest green to rusty brown, usually light green, scrambling over twiggy branches and shrubs and moss cushions. Stems to 15 cm long, 1–2 pinnately branched pinnae and pinnules of varying lengths, pinnae often a little deflexed, may be short and regular in length. Leaves of the stem, 1.4 mm wide, plano-distichous or sub-amplexicaul with apices deflexed, apiculate, the bases of both margins laciniate-toothed in varying degrees, and sometimes ciliate towards the apex; lobule a spiniferous lamina, variable in form, or with a lamina interposed between the stem and a saccate lobule, which may itself bear a spine from the fissure at the side; leaves of the branches obliquely ovate, ciliate-toothed all round, often spinous at the bases; lobules clavate-saccate, usually with a small adjoining spiniferous lamina or stylus. Underleaves of the stem, 4-fid, all irregularly spinose-dentate, of the branches, 2–4, usually 4-fid, margins of segments less spinous toothed than those of the stem, 2 median segments sometimes clavate-saccate; of the branchlets, may be bis-bifid, segments plane or saccate. Cells ca. 25μ, round or oval, trigones present. ♀ inflorescence terminal on the stem as though in the fork of two apical pinnae, ovate-oblong; floral leaves and underleaves irregularly elongated, laciniate-toothed, merging into spinous-dentate paraphylls sparsely or densely clothing the coelocaule containing the sporophyte, with the apex crowned with sterile pistillidia carried upwards on the growing perigynium. ♂ bracts terminal on pinnae and pinnulae, in 3–4 pairs, crowded, concave, conduplicate, lobule smallish, apex of lobe trispinose, antheridia rarely present.
L. palpebrifolia whose name means leaves fringed as with eyelashes, is recognized by its ciliate-toothed branched leaves, which are not much smaller than the stem
leaves, and its usually pale colour and soft texture. Both dorsal and ventral bases of the cauline leaves are laciniate-toothed, and some are toothed towards the apex. Specimens in which the leaves are flat and not amplexicaul have a different appearance from those in which the leaves loosely, or closely, clasp the stem. These forms drift into one another. It is nearest to L. Taylori, from which it differs in the ciliate-toothed branch, and partly ciliated stem leaves. L. Menzies has longer marginal cilia and smaller branch leaves. From L. clavigera it is differentiated by its larger size, spinous-toothed underleaves, and the toothed paraphylls on the coelocaule when this is present.
North Island: Waipoua Forest, H782, K. W. A.; Mangatihi, W. slopes of Mt. Ruapehu, 18933, Auckland Museum Herb., J. E. Atwood; Waikare-iti Urewera, 3,000ft (red-brown), 10595, E. A. H; Huirau, Urewera, 3,200ft, 10,933ft, H. M. D.; Upper Ruakituri R., 1,500ft, 10392, B. Teague; Pohangina V., 442 A. P. D.; South Rimutaka Ranges, 5783, H. H M.; Eastbourne, 2889, N. J. Butler; Mangaroa, 4216, S. Berggren; a1751 Herb. Colenso; Whakapapa, National Park, 141, G. O. K. S.
South Island: Marlborough, 88, 285, J. H. McMahon; Nelson, Kaituna Creek, 5807, G. S.; Pelorous Bridge, 6317, I. Haskell; Arthur's Pass, 5858, Mrs. Vere, 1, 2, W. M.; Caswell Sound, 6330, V. D. Z.; Waiho, 10756, W. M.
The type was collected at Dusky Sound by Dr. Archibald Menzies. Endemic as far as is known.
Lepidolaena Taylori (Gottsche) Trevisan.
Polyotus Taylori G. in G.L. et N., Syn. Hep., 246, 1845.
Polyotus claviger var. Taylori Mitt. in Fl. Nov. Zel., ii, 152, 1854; Hook. Handb. N.Z. Fl., ii, 529, 1867.
Lepidolaena Taylori Trev. Sch. di. una class. delle epatiche, 393, 1877; Steph., Spec. Hep., iv, 47, 1909.
Polyotus fimbriatus Col., Trans. N.Z. Inst., 19, 294, 1886.
Polyotus smaragdinus Col., Trans. N.Z. Inst., 19, 294, 1886.
Polyotus prehensilis Col. Trans. N.Z. Inst., 21, 69, 1888.
Plants dioicous, robust, on trees and shrubs, dull green to red-brown, variable. Stem to 15 cm, regularly to irregularly bi-tripinnate, pinnae short or long, usually decurved when short, 3–5 mm, interspersed with longer ones which may be again bipinnate. Leaves, of the stem, contiguous to imbricate, appressed, may be squarrose if crowded, diverging somewhat, broadly ovate, to 1.8 mm across, lacinate-toothed at the bases of the leaves, more particularly the dorsal base, acute to mucronate; lobules bifid, segments variously shaped, consisting of a spinose triangular lamina, and a smaller convolute segment; leaves of the branches, closely imbricate, smaller, occasionally toothed towards the apex, lobule clavate-saccate, with 1 or 2 spines, with a small intervening triangular lamina. Underleaves, of the stem, distant, 4-fid, margins spinous-toothed; of the branches and branchlets, 4-fid with the 2 median segments saccate with or without 2 short spines, but sometimes reduced to only 2 segments, saccate or plane of 1 of each. Cell cavities rounded, oval or irregular, 30μ with both larger and smaller intermixed. ♀ inflorescence usually terminal with 1 subfloral innovation; floral leaves and under-leaves more spinous-toothed than in L. clavigera; coelocaule obovate, walls 6–8 cells thick, densely paraphyllous, paraphylls with toothed margins, very irregular in size and shape, sporophyte not seen. ♂ bracts terminal on the pinnulae (branchlets), in 2–3 pairs, complicate, lobule large, apex free, acute to mucronate.
This is the commonest of the New Zealand species. It is a larger plant than L. claviger, is never glaucous, has spinous-toothed margins to the underleaves and more spines about the lobules. Mitten's having reduced it to a variety of L. claviger (1854) has caused much confusion. Simple forms of L. Taylori may have the laciniate leaf-bases showing as simply (but perhaps longly) dentate, as in Gottsche's description.
A specimen from Caswell Sound, 10981, V. D. Z., is intermediate in form between L. Taylori and L. Berggrenii. The underleaves in some cases are ciliate on the margins, and the ventral leaf-bases with 3 longish teeth sometimes present as in the type, but the branch leaves are small and toothed. A somewhat similar
Explanation of Text-figure
St leaf—stem leaf; br. leaf—branch leaf; st. underlf.—stem underleaf; br underlf.—branch underleaf, fl. leaf—floral leaf; fl. underlf.—floral underleaf.
Figs. 1–7.—L. clavigera Fig. 1—st. leaf. Fig. 2—fl. leaf. Fig. 3—coclocaule. Fig. 4—st. underlf. Fig. 5—X section of seta. Fig. 6—br underlf. Fig. 7—fl. underlf. Figs. 8–12.—L. palpebrifolia.. Fig. 8—Juvenile inflorescence showing archegonia. Fig. 9—st. lvs. Fig. 10—fl. underlf. Fig. 11—st. underlf. Fig. 12—st. leaf. Figs. 13–18—L. Taylori. Fig. 13—st. lvs. Fig. 14—fl. lvs. Fig. 15—fl. underlf. Fig. 16—fl. leaf. Fig. 17—Minute growths from germinating spores among the paraphylls. Fig. 18—st. underlf. Figs. 19–25—L. Berggrenu. Fig. 19—branchlet. Fig. 20—st. leaf. Fig. 21—Paraphylls of coclocaule. Fig. 22—st. lvs. Fig. 23—st. underlf. Fig. 24—♂ bracts. Fig. 25—fl. underlf. Figs. 26–27—L. allophylla. Fig. 26—branches ventral. Fig. 27—stem, ventral Figs. 28–30—L. reticulata. Fig. 28—st. lvs. and underlvs. Fig. 29—br. leaf. Fig. 30—br. underlvs Figs. 31–36—L. Menziesn. Fig. 31—fl. leaf. Fig. 32—coelocaule. Fig. 33—st. leaf. Fig. 34—st. underlf. Fig. 35—br. underlf. Fig. 36—br. leaf. Figs. 37–46—L. Magellanica. Fig. 37—coelocaule. Fig. 38—fl. leaf. Fig. 39—st. underlvs. Fig. 40—leaf. and lobule. Fig. 41—branch, ventral.
specimen is from Marlborough. While specimens of L. reticulata from Antipodes, from the dorsal aspect, are indistinguishable from small stems of L. Taylori.
Colenso's P. prehensilis, named from the prehensile-looking branches, is a graceful, dull green plant with many longish branches, a form often seen. This, as well as P. fimbriatus were published as synonyms by Stephani (1892). P. smaragdinus Col., “a pleasing emerald green,” has all the characteristics of L. Taylori.
Plants examined include at least 15 from Berggren's collection of 1874. Also from the Chatham Islands, 9593, W. Martin, 1925.
The type was from Dusky Sound, coll. Menzies. Endemic to New Zealand as far as is known.
Lepidolaena reticulata (Tayl.) Trev.
Jungermannia reticulata Tayl., Lond. Journ. of Bot., 395, 1844.
Polyotus reticulatus Gottsche in G. L. et N., Syn. Hep., 245, 1845; Mitt. Handb. N.Z. Fl., ii, 755 (Appendix) 1876.
Frullania reticulata Hook. Handb. N.Z. Fl., ii, 531, 1867.
Lepidolaena reticulata (Tayl.) Trev. Sch. di una class delle epatiche, 393, 1877; Steph., Spec. Hep., iv, 46, 1909.
Plants dioicous, small to medium, on and amongst other bryophytes, brown to reddish brown, or purplish to blackish. Stem to 3 cm, flexuous, moderately to closely bipinnate, pinnae to 1 cm, often crowded, rarely tripinnate. Leaves of the stem, usually imbricate, obliquely ovate, ca. 0.8 mm, entire or with 1 or 2 teeth at the ventral base, decurved, apex rounded or cuspidate, lobules bifid, 1 segment clavate-saccate, the other narrowly triangular, or greatly reduced or merely a spine; leaves of the branches (pinnae) obliquely rounded-ovate, upper ones occasionally a little toothed, lobules saccate, erect, or deflexed, appearing pedicillate. Underleaves of the stem variable, usually 4-fid to ½ or ⅔, with segments narrowly triangular or lanceolate, margins not reflexed, entire; of the branches, usually bifid, but on the primary branches may be 3–4 fid, from a cellular base 4–6 cells deep, segments plane, usually tapering, otherwise 2 clavate segments from a cellular base. Cells rarely more than 30μ, except at the base of the leaf; cortical cells quadrate to rectangular, 20–30μ wide. ♂ bracts terminal on the pinnules in 2–3 pairs, apex usually free with an apical tooth. ♀ inflorescence not seen.
L. reticulata differs from L. clavigera in the plane margins of the segments of the stem underleaves. In L. clavigera these margins are reflexed and the lobules crenulate with convex cell-walls. Other points noticed about L. reticulata are, that when the stem underleaves have 2 saccate segments, these are on the outside with the 2 middle ones plane, whereas in L clavigera, if there are 2 saccate segments of the underleaves, these are in the middle, with the 2 non-saccate ones on the outsides; on some stems there are at the bases of the underleaves, small brown minutely cellular laminae, each bearing a tuft of rhizoids, but this is probably to be found in other species also.
Typical specimens of L. reticulata do not appear to be common on the 2 main islands of New Zealand, but there appear to be intermediates between this species and L. Taylori.
The specimens from the Antipodes have short outgrowths, probably incipient spines, on the lobules, and occasionally the stem underleaves are spurred.
South Island: Rocky bank with Chandonanthus squarrosus, first stream west of Arthur's Pass township, 2,600ft, 8, W. M; Revolver Cove, Preservation Inlet, Fiordland, 5844, H.H.A., 1946; near Tin Range, Stewart Island, 11145, W. M.
Antipodes Islands, on the ground in tussock country, 2597, packets 4, 5, 6, G. Einar Du Rietz, April, 1927.
Auckland Island, Musgrave Peninsula, climbing amongst Lepicolea scolopendra, 10972, R. L. Oliver, January, 1944.
The type was from Auckland Island, coll. Hooker. Stephani also reports it from Desolation Island.
Lepidolaena Berggrenii Hodgson
Planta dioica, mediocris, corticola, flavo- vel rufo-brunnea. Caulis procumbens, 4–5 cm longus, repens, regulariter bipinnatus, pinnis remotiusculis, ca. 4 mm longis, pinnulis vulgare
quadrijugis, ca. 2 mm longis. Folia caulina nitida, remota vel contigua, 0.7 mm longa et lata, oblique cordata, obtusa, margine ventrali inferne brevi vel spinoso-dentato; lobulo evoluto vel clavato-saccato, lamina interposita lata et integerrima vel varie biloba. Cellulae incrassatae, plerumque rotundae, aliis ovalibus, aut angulatis, ca. 30μ, trigonis parvis. Folia ramea minora, 0.5 mm, ovata, acuta ad rarius aristata vel obtusa, integerrima vel angulata vel dentata. Folia ramulina minuta, angusta, 0.3–0.4 mm × 0.2 mm longa dentata, lobulis clavatis, a caule remotis, pedicellatis. Amphigastria caulina ad 0.4 mm, quadrifida, segmentis e basi lata, planis, obtusis vel attenuatis, integerrimis; ramorum quadrifida, medianis segmentis planis, externis clavato-saccatis; ramulina bi-saccata e lamina basali. Gynaecium terminale, foliis involucralibus majoribus, ovatis, acutis vel mucronatis, recurvo-concavis; lobulis variis; amphigastria quadrifida, segmentis aliquantum dentatis, elongatis; coelocaule vetus 2.5 mm pyriforme, paraphyllia superne oblonga vel anguste ovata, integerrima, inferne subnulla (in typo). Androecia terminalia in pinnis, bracteis ad 5-jugis, conduplicatis, breve bilobis.
Plants dioicous, medium, greenish to reddish brown, corticolous, in mountainous country. Stem to 5 cm long but often shorter, creeping on bark, regularly bipinnately branched, pinnae to 4 mm, pinnules ca. 2 mm, about 4 pairs to each pinna, cortical cells 2–3 times longer than broad. Leaves of the stem glossy, remote to contiguous or a little imbricate, to 0.75 mm long and broad, obliquely cordate with an arched insertion ventral margin variously toothed to about half way or less, teeth short or merely unevenesses of the margin, except in the basal part; lobules clavate-saccate or evolute, with an intervening lamina which may be broad and entire, or variously bilobed. Cells averaging ca. 30μ, mostly rounded, or oval and longer, or even angled, trigones usually small. Leaves of the branches smaller, 0.5 mm ovate, acute to rarely aristate, apiculate or obtuse, entire or angled or a little toothed. Leaves of the branchlets (pinnulae) minute, narrow, 0.35–0.4 mm by 0.2 mm, longly toothed, lobules distant from the stem and appearing stalked, usually simply clavate-saccate. Underleaves of the stem to 0.4 mm, 4-fid, segments flat, tapering or obtuse, 6–8 cells broad at the base, margins entire; of the branches, 4-fid with the 2 median segments plane, the outside ones saccate; of the branchlets, 2-fid, both segments saccate, from a basal lamina. ♀ Inflorescence terminal on the main stem, in the fork of an innovation from among the floral leaves and a branch arising from a little lower down, in the early stage with a tuft of archegonia at the extremity of the stem surrounded with floral leaves. Floral leaves numerous, variable, narrower and longer than the cauline, sparsely toothed, plane or recurved and concave, lobules not saccate, irregular in shape. Underleaves large, deeply 4-fid, segments a little toothed. Coelocaule pyriform, 2.5 mm, vested with paraphylls near the mouth on one old one. Others may have the paraphylls evenly distributed. Paraphylls oblong to narrow ovate, entire, cells large, hyaline. Androecia in 2–3 pairs at the terminus of a primary branch. Bracts inflated, imbricate, shortly and subequally bilobed, antheridia 2 mm in diameter, underleaves bilobed, segments plane.
This species is nearest to L. reticulata, from which it differs in the glossy leaf-surface, in the minute, narrow, toothed leaves of the pinnules, and in the habit and habitat, creeping on bark, mostly of scrub, often with Frullanias. With underleaves and lobules similar to those of L. reticulata, it bears the same relationship to this species as does L. palpebrifolia to L. Taylori. It is probably not uncommon in New Zealand, and is easily recognized by its shiny and prickly appearance.
North Island: On Dracophyllum sp. Mangawaru Plateau, Raukumara Ranges, approx. 4,300ft, 9288, Mt. Wharite, S. Ruahines, 2,500ft, 5787, A.P.D.; Otupae, N.W. Ruahines, 2–3,000ft, 1591, (Type), 3672, A. P. & H. M. D.; Ruahines summit, Herb. Mitten, 2078, Colenso, sub. nom. L. Brachyclada; Mt. Egmont., 5819, G. O. K. S., 10341, E. A. H.; Mt. Alpha, Tararuas, 3,900ft, 2461, A. P. D.
South Island: Otira, 3447, in alpibus ad flumen Otira, 3444, S. Berggren, 1874; Misery Creek, Cass, North Canterbury, B27, L. Visch, comm. W. R. Philipson; also H5, Cass.
The type is 1591 from Otupae, N. W. Ruahine Ranges, on Leptospermum scoparium near bog, H. M. Druce. It is named for Dr. Sven Berggren, a veteran collector of New Zealand hepatics, though Colenso's gathering from the summit of the Ruahine Ranges is recorded 20 years earlier under the name of P. brachyclada in Flora Novae Zelandiae ii, p. 152, 1854.
Lepidolaena allophylla (Tayl.) Trev.
Jungermannia allophylla Tayl. Lond. Journ. of Bot., 395, 1844.
Polyotus allophyllus Gottsche in G. L. et N., Syn. Hep., 249, 1845; Mitt. Appen. Handb. N.Z. Fl., ii, 755, 1876.
Frullania allophylla Hook. Handb. N.Z. Fl., ii, 538, 1867.
Lepidolaena allophylla (Tayl.) Trev. Sch. di una class. delle epatiche, 393 1877; Steph. Spec. Hep., iv, 46, 1909.
Plants sterile, very small, reddish to crimson-purple. Stems barely 1 cm, 0.2 mm wide, flexuous, almost geniculate, regularly pinnately branched; leaves of the stem contiguous, to 0.5 mm, broadly oval, entire or sparsely toothed, lobules clavate-saccate, sometimes with a minute stylus, 0.2 mm long, 0.1 mm wide at the rounded apex, appearing pedicillate from being joined with the incurved margin of the stem, deflexed to horizontal; leaves of the branches smaller to 0.4 mm, variable in shape, 2–3-toothed to spinous-toothed near the apex; lobules as on the main stems. Underleaves of the stem ovate to rotund or broadly ovate, bilobed-emarginate, as in Frullania, sinus obtuse, gaping, lobes obtuse or acute, margins entire; of the branches, bifid, lobes usually clavate-saccate. Cells ca. 30μ, which is large for such a small plant, quadrate-hexagonal, walls thickish, trigones minute.
Although I cannot account for the fact that the fragmentary piece of “type” of L. allophylla in Mitten's Herbarium is undoubtedly L. reticulata, with 4-fid stem underleaves, nevertheless there definitely is this minute species in the subantarctic islands, answering to Taylor's description of his J. allophylla, and distinct from L. reticulata in its small size and bilobed stem underleaves, and the spinous teeth on the branches. The lobules at times also, are more deflexed, sometimes lying horizontal, as in Taylor's illustration.
Lepidolaena Hariotana (B. & M.) Schiff. from Tierra del Fuego, 133 P. Dusen, 1896, also has rounded bilobed stem underleaves, and differs only in its larger size and entire branch leaves.
Port Ross, Auckland Islands, in cushion of Gaimardia ciliata, 2282, G. Einar Du Rietz, 1927.
Campbell Island, with other hepatics on lichen in scrub, North-East Harbour, 2578, G. Einar Du Rietz, 1927.
The type was from Campbell Island, on hills, mixed with other hepatics, coll. Hooker, whose comment was, a very curious and beautiful little plant.
Lepidolaena Menziesii (Hook.) Trev. ex Dumort.
Jungermannia Menziesii Hook. senr., Musci Exotict ii, tab. 118, 1820.
Lepidolaena Menziesii Dumort. Rec. d'Obs. sur les Jung., 13, 1835 nom. nud.
Trev., Sch. di una nuova class. delle epat., 393, 1877; Steph., Spec. Hep. iv, 50, 1909.
Polyotus Menziesii Gottsche in G. L. et N., Syn. Hep. 247, 1845.
Plants dioicous, pale green when fresh, fading to whitish or light brown, creeping on earth and humus, mainly on beech forest floor and bogs (in New Zealand), everywhere ciliate. Stems to 6 cm. (Stephani gives 10), simple, often forked at the apex, closely and evenly bipinnate, pinnae ca. 5–15 mm long, usually ca. 8 mm shortening towards the apex of the stem as in a fern frond, horizontally spreading, sparingly to densely paraphyllous, paraphylls simply ciliform to 4–5 ciliate from a variously shaped base. Cauline leaves 1 mm, including the cilia, distant or more rarely imbricate, ovate-cordate, longly ciliate, cilia articulated, to as many as 5 cells in length, lobules concave, margins ciliate; of the pinnae similar but smaller, a little imbricate, of the pinnulae still smaller, more vertically set and in 2 distinct rows; lobules saccate with an interposed lamina. Underleaves of the stem deeply 4-fid, margins recurved, and copiously and longly ciliated; of the pinnae and pinnulae 4-fid, with the 2 median segments saccate. Cell cavities ca. 20μ, basal a little longer, trigones large. ♀ inflorescence terminal in the fork of 2 sub-floral branches; floral leaves narrow ovate-triangular, longly ciliate, underleaves 4-fid, laciniate, coelocaule elongated, 1.2 mm wide, lengthening from the stem apex to as much as 8 mm, fleshy, tubular, wall ca. 8 cells thick, enveloping a solid structure which is the growing seta of the capsule. This growth keeps pace with that of the coelocaule (stem perianth) as vertical sections at different stages showed the sporophyte completely filling the coelocaule. The floral leaves and underleaves may be carried upwards with the growth of the coelocaule, and are hard to distinguish from the paraphylls. The coelocaule widens somewhat towards the piliferous apex. Exserted capsule and ♂ inflorescence not seen.
Although lacking any roseate or purplish colouration, L. Menziesii is nevertheless a very beautiful species. It is easily recognized by its pale colouring, densely bi-pinnate and paraphyllous stems, and all leaves very longly ciliate. Beckett was probably the first to discover the species in New Zealand, from Teremakau, Westland, Nos. 386 and 4287 Herb. Levier, 1887 and 1903 respectively, also from Alford forest Canterbury, 387, 1893. Fresh moist specimens stain the paper green or greenish yellow.
North Island: Mt. Ruapehu 351, Mt. Egmont 5799, G. O. K. S.; National Park, H165, L. B. M.; gorge above Rangitikei Forks 3617; bog, Aniwaniwa V. Urewera, 9315, H. M. D.; N.W. Ruahines, 3533, bog on Mt. Egmont, 3647, 3,000ft, Mangaroa 698, bog above bush, Tararuas Mts, A. P. D.; Ohau R., P.R.B. Herb., V. D. Z.
South Island: Maitai V. Nelson, 5971, B. M. Kidson; Arthur's Pass, 5800, Mrs. Vere; Arthur's Pass, 24, 48, 10776, L. Te Anau, 5613, L. Manapouri, 5615, W. M.; Torlesse Range, 5828, H. M. H.; Castle Hill, 437, S. Berggren; Fox Glacier, 5806, Mrs. Knight; Milford Track, 1618, F. Macdonald, 5806, Miss Bevan; Haast Pass, H611, M. P. Matthews, H5965, K. W. A. (fruiting); Wilkin R., L. Wanaka 5796, L. Manapouri, H831, Freeman R., 36657, P.R.B. Herb., G. S.; Conical Hill, Otago, 17158, P.R.B. Herb., G. B. Rowlings; Cass Hut, North Canterbury 5801 (fertile) E. Ensor.
Campbell Island: Tucker Cove, 5882, R. L. Oliver, 9/10/44; J. H. Sorensen (fruiting) 5870, 4/1946; open scrub, 26 W. B. Brockie (fruiting) 4/1947.
The type is given as from Staten Land, near Cape Horn, coll. Menzies. Also from South America.
Lepidolaena magellanica (Lamarck) Schiff.
Jungermannia magellanica Lamarck Encycl. Bot. (Hepaticae) iii, 28, 1791, Hook. senr. Musci Exotici ii, tab. 115, 1820.
Polyotus magellanicus G. in G.L. et N., Syn. Hep., 248, 1845; Mitt. in Hook. Fl. Tas. 1860; Hook. Handb. N.Z. Fl., ii, 529, 1867.
Lepidolaena magellanica (Lam.) Schiff. in Engler-Prantl. Natürl. Pf. Fam. Hepaticae, 1893; Steph. Spec. Hep., iv, 48, 1909; Rod. Pps. & Trans. Roy. Soc. Tas. Bry., ii, 1916.
Plants diocious, robust, forming compact or straggly mats on earth or logs, brown to brownish red. Stems short or long, 2–15 cm, according to habitat and environment, mostly simple, closely or loosely bi-tripinnate, pinnae horizontal, deflexed or circinate when dry, very regular, length ca. 1 cm, but varying in proportion to the length of the stem. Leaves, of the stem, imbricate to 1 mm long, obliquely broadly ovate or cordate, upper margin (dorsal) often reflexed, margins ciliate, specially towards the base; lobule bifid, one segment saccate with or without a spine, the other a triangular lamina usually spinous, generally hidden by the broad underleaves; leaves of the pinnae (primary branches) somewhat smaller, spinous-ciliate, lobule clavate-saccate, distant and deflexed from the stem, mostly with one spine, or a volute; of the pinnulae (ultimate branches) much smaller, ciliate, lobules distant from the stem and deflexed, generally with a spine. Underleaves of the stem large, reniform, obscuring the leaf-lobules, to orbicular-ovate emarginate-bifid, segments acute to mucronate, margins entire to sparsely ciliate, reflexed, of the pinnae, more deeply incosed, ovate, spinous, of the pinnulae similar, or often with 2 saccate segments Cells variable, in some plants porose and small ca. 20μ or less with very thick walls, in others rounded-oval to 30μ or more, trigones large. ♀ inflorescence lateral ca. 6 mm, floral leaves and underleaves in several rows at the base, elongated, imbricate, bifid, the shorter segment doubtless representing the lobule, concave enveloping the coelocaule almost to the apex apices and margins longly piliferous, hairs much tangled. Coelocaule narrow-oblong, glabrous smooth, flimsy, mouth piliferous; at least 10 archegonia were counted in the midst of the floral leaves, before the coelocaule had begun to form; capsule 2.8 mm, valves cohering at the apex in one specimen with lateral margins reflexed, seta 6 mm, no spores seen ♂ bracts terminal in few pairs on the branches, bilobed, conduplicate, imbricate, entire, with an apical spine, on the dorsal lobe.
In explaining the differences between L. magellanica and L. Ljungneri from Patagonia, and an undescribed Tasmanian species named L. Weindoerferi, Dr.
Herzog (1940, p. 8) states that these 2 species are very similar to L. magellanica, but that the latter differs in its reniform, undivided, unarmed stem underleaves, as represented in the accompanying drawing. This is not correct, as there is always a sinus at the apex, though this may be hard to detect against the stem, and all descriptions agree on this point Also the margins may be ciliate. To quote the Synopsis Hepaticarum: “… amphigastriis caulinis orbiculato-ovatis, emarginatobifidis, integerrimis ciliatisve, …” Thus, the other mentioned distinction between L. magellanica and L. Ljungneri breaks down.
This species can be readily recognized by its brown to reddish colour, and the large reniform stem underleaves, in conjunction with the spinous-ciliate branch leaves.
North Island: Wairoa R. Kaipara, 789 Mossman; forming mat on summit rocks with Hymenophyllum, Te Moehau Mountain, 20, Mt. Alpha, Tararua Ranges in beech forest, H160, L. B. M.; on logs with Dicranoloma sp. western shore of L. Waikare-iti, 3,000ft; Urewera National Park, 5833, E. A. H.; Tararuas, 5830, A. D. Beddie; Wairarapa Forests, Colenso.
South Island: Westland, 243, T. Kirk; Milford Track, 1627, F. Macdonald; Caswell Sound, V. D. Z.; Doubtful Sound, 6700, W. M.; Freeman R., L. Manapouri, 36659, P.R.B. Herb., G. S.; Routeburn V., 1824, G. Einar Du Rietz; on log, to L. Manapouri, 5631, W. M.
Stewart Island: 267, T. Kirk; wet ground, forest clearing, Tin Range, 509, ground in manuka, Crooked Reach, 656, W. M.
Campbell Island, on alpine rocks, J. D. Hooker (Handbook); also from Tasmania, Australia, Cunningham, and near Cape Horn, Menzies, 1787.
The type was from the Straits of Magellan, coll. Commerson.
The Kaipara specimen coll. Mossman, 1850, may have been the first of this species to be collected in New Zealand, though Colenso's gatherings are recorded in the Flora Novae Zelandiae ii, 1854.
Lepidolaena brachyclada (Lehm.) Trev. is a Tasmanian and Australian species, which in my opinion does not grow in New Zealand. It is a robust species with very long stems, and short pinnate branches as the name implies. The stem as well as the branch leaves are ciliated all round, the cilia to as long as 12 cells, and widely spaced. The lobules are thin and often bi-lipped at the mouth, and the stem underleaves are more or less armed with slender spines. The specimen in Mitten's herbarium, No. 2078 Colenso, identified by Mitten as Polyotus brachycladus has none of these characteristics, and is L. Berggrenii sp. nov. In this specimen the stem leaves are toothed only at the ventral base and the lower ventral marign, the ultimate branch leaves being longly toothed as is characteristic in this newly described species It was probably these longly toothed branch leaves which misled Mitten as to its identification.
It was collected in Tasmania in 1824 by Neill and described in Pugillus viii by Lehmann as Frullania brachyclada, also collected in Australia by Fraser. It now appears to be somewhat rare.
The 3 remaining species of the genus (not already mentioned) are, as named and described in Stephani's Species Hepaticarum vol. vi.
L. Halliana, a large species with 4-fid underleaves from Patagonia.
L. Patagonica, small with reniform bifid underleaves (apparently similar to L. allophylla), from Southern Patagonia.
L. Skottsbergii St., large with reniform bilobed underleaves from Falkland Islands.
My grateful thanks are due to Dr. C. T. Rogerson, of the New York Herbarium, for the loan of specimens from Mitten's collection, to Dr. C. E. B. Bonner, of Geneva, for information and the loan of specimens, to Dr. J. Proskauer, of the University of California, for the gift of a copy of Dumortier's “Recueil”, to Mr. V. D. Zotov, of the Botany Division, D.S.I.R., Christchurch, for advice on nomenclature, to Dr. Tycho Norlindh, of the University of Lund, Dr. H. Persson, of the Riksmuseum, Stockholm, Dr. H. N. Barber, of the University of Tasmania, and Mr. R. T. M. Pescott, of the Melbourne National Herbarium, for material assistance, and a special thank you to Sir Edward Salisbury late of Kew, for the loan of the original specimen of L. clavigera.
Colenso, Rev. Wm., 1886. A Description of some newly discovered cryptogamic plants. Trans. N.Z. Inst., 19, 294, Wellington.
— 1888. Trans. N.Z. Inst., 21, 68 Wellington.
Dumortier, B. C., 1835. Recueil d'Observations sur les Jungermanniacees, Tournay.
Gottsche, C. M., 1845. In G. L. et N., Synopsis Hepaticarum, pp. 244–249, Hamburg.
Hatcher, R. E., 1958. The Genus Trichocolea in N.Z., Trans. Roy. Soc. N.Z. 85, 237–246.
Herzog, Dr. Th., 1940. Die Moose der Exped. Ljungner nach Patagonien, Arkiv för Botanik Band 29 A. No. 21, Stockholm.
Hodgson, E. A., 1950. In Martin W. Bryophytes of Stewart Is ii, Trans. Roy. Soc. N.Z. 78, 486–581.
Hooker, Sir J. D., 1867. Handbook of the N.Z. Flora, ii, London.
— 1818–1820. Musci Exotici i and ii, London.
*Lamarck, J B. A. de, 1791. Tableau encyclopedique et methodique des 3 regnes de la Nature (Hepaticae), 476–480.
Lindberg, Dr. S. O., 1872. On Zoopsis, Journ. Linn. Soc. Bot., 13, 190 footnote, London.
Mitten, W., 1854–1855 Fl. Nov. Zel., 11, London.
— 1860. Fl. Tas., London.
Pearson, W. H., 1923. Notes on a Collection of N.Z. Hepatics, Univ. Cal. Pub. Bot. 10.
Rodway, L., 1916. Tas. Bry ii, Hepatics, Pps. & Proc. Roy. Soc. Tas., Hobart.
*Schiffner, V., 1893. Hepaticae in Engler-Prantl. Naturl. Pflanzenfamilien, Leipzig.
Schuster, R. M., 1858. Annotated key to the orders, families, and genera of Hepatics of America north of Mexico, The Bryologist 61, 1–66, Durham, N.C.
Stephani, Dr. Fr., 1909 and 1924. Species Hepaticarum iv and vi, Geneva.
— 1892. Colenso's N.Z. Hepaticae, Journ. Linn. Soc. Bot. 29, 263–280.
*Trevisan Di Saint Leon V., 1877. Schema di una nuova class. delle Epatiche, Milan.
Mrs. E. A. Hodgson,F.L.S.,
Kiwi Valley R.D.,
Wairoa, Hawke's Bay.
[Footnote] * Original not seen.