Tuatara: Volume 3, Issue 1, May 1950

The Classification of New Zealand Hepaticae

The Classification of New Zealand Hepaticae

The Hepaticae or liverworts form the second of the two classes into which the Bryophyta are divided. The two main distinctions between the classes are said to be, that of the liverworts have unicellular rhizoids, and spiral elaters mixed with the spores in the capsules (except in Ricciaceae). The non-thalloid or foliose hepatics differ from the mosses, generally speaking, in the following respects; in the rudimentary and short-lived protonema; in the bi-lateral leaves always without a midrib, and frequently bilobed; in the usual presence of stipules; in the capsule remaining in the more or less delicate calyptra (destitute of lid) till the spores are mature, then developing a hyaline and evanescent, sometimes elongated seta, and commonly dehiscing by four valves, peristome always absent. Generally speaking, the liverworts prefer wetter conditions than the mosses.

Probably the earliest collection of New Zealand hepatics was that of Archibald Menzies, gathered at Dusky Sound in 1791. These were described and figured by Sir William Hooker in his Musci Exotici in 1818. Sir Joseph Hooker's Botany of the Antarctic Voyage, in several volumes spread over many years, contains the descriptions of plants collected by Hooker in the subantarctic islands, Northland (N.Z.), and Tasmania, together with many of Colenso's. Dr. T. Taylor and W. Mitten were responsible for these descriptions. In 1867, Part II of Hooker's Handbook of the New Zealand Flora was published, a portion of which is devoted to the liverworts, with an Appendix by Mitten. Dealing with plants collected by the earlier botanists, the Handbook has been the basis of all our work on New Zealand Hepaticae. Stephani's famous Species Hepaticarum in 6 volumes, 1900-1924, includes descriptions of all New Zealand species to date. Local Floras of both hemispheres are useful for obtaining information about hepatics, the handiest, perhaps, being L. Rodway's “Tasmanian Bryophyta II, Hepatics.” *

The system of classification used in this paper, is that set out by Evans. **

Three of the four orders of the Hepaticae are represented in New Zealand.

* Papers & Proc. Roy. Soc. Tasmania 1916.

** Evans, A. W., 1939. Classification of the Hepaticae, Botanical Review, vol. 5, p. 20.

Order 1 Jungermanniales

Plants both foliose and thalloid, the latter not differentiated into layers of different tissue and without pores. Tuberculate rhizoids not present. Sexual organs usually in groups, but not on special pedunculate receptacles. Capsule, usually on a long seta, opening by 4 valves. Apical growth of stem or thallus proceeds from a single cell.

Sub-order 1 Haplomitrineae

  • Family Haplomitriaceae, Calobryum.

Sub-order 2 Jungermannineae (Jungermanniales acrogynae)

  • Family Ptilidiaceae, Herberta, Mastigophora, Hygrobiella, Ptilidium, Bleoharostoma, Isotachis, Lepicolea, Lepidolaena, Trichocolea.
  • Family Lepidoziaceae, Bazzania. Acromastigum, Psiloclada, Lepidozia.
  • Family Cephaloziaceae, Cephalozia, Lembidium, Zoopsis, Adelanthus, Marsupidium.
  • Family Cephaloziellaceae, Cephaloziella.
  • Family Harpanthaceae, Lophocolea, Chiloscyphus, Mylia, Geocalyx, Saccogyna.
  • Family Jungermanniaceae, Lophozia, Sphenolobus, Anastrophyllum, Cuspidatula, Chandonathus, Jungermannia, Jamesoniella, Acrobolbus, Symphyomitra.
  • Family Marsupellaceae, Gymnomitrium.
  • Family Plagiochilaceae, Plagiochila, Tylimanthus.
  • Family Scapaniaceae, Diplophyllum.
  • Family Schistochilaceae, Schistochila, Balantiopsis.
  • Family Porellaceae, Porella.
  • Family Goebeliellaceae, Goebeliella.
  • Family Radulaceae, Radula.
  • Family Frullaniceae, Frullania.
  • Family Lejeuneaceae, 18 genera, originally subgenera of Lejeunea, keyed separately.

Sub-order 3 Metzgerineae (Jungermanniales anacrogynae)

  • Family Treubiaceae, Treubia.
  • Family Fossombroniaceae, Fossombronia Petalophyllum.
  • Family Pelliaceae, Calycularia, Allisonia.
  • Family Pallavicineaceae, Pallavicinia, Symphyogyna, Hymenophytum.
  • Family Metzgericeae, Metzgeria.
  • Family Riccardiaceae, Riccardia.
  • Family Monocleaceae, Monoclea.

Order 3 Anthocerotales

Thalloid, with smooth rhizoids, no ventral scales. Cells usually with one large chloroplast. Sporogonium with a bulbous foot, a sheath, and a long sessile capsule, bursting from the top downwards into two valves, columella generally present. Antheridia sunk in the upper surface of the thallus, ultimately bursting free.

Family Anthocerotaceae, Anthoceros, Aspiromitus (on the authority of Pearson), Megaceros, Dendroceros.

Key to Genera

1. Plants foliose 2
Plants thalloid 43
2. Leaves (plus or minus) reduced to cauline lobes, cells inflated Zoopsis
Leaves not reduced to cauline lobes 3
3 Leaves spirally arranged Calobryum
Leaves not spirally arranged 4
4. Archegonia on the upper surface of a thalloid stem (Fig. 30, P) Fossombronia
Archegonia at first terminal on stem or branch, involucre formed from true leaves 5
5. Elaters with one spiral band, capsule valves not split to the base 6
Elaters with 2 or more spiral bands, capsule valves split to the base 7
6. Branches intra-axillary, lobule convex towards the upper lobe (Fig. 13, B) Frullania
Branches infra-axillary, lobule concave towards the upper lobe Lejeuneaceae
7. Leaves incubous or transverse 8
Leaves succubous to transverse 19
8. Leaves with an upper (dorsal) lobe and lower (ventral) lobule 9
Leaves without a ventral lobule 12
9. Dorsal lobe and ventral lobule closely complicate 10
Lobe and lobule not closely complicate 11
10. Stipules present Porella
Stipules absent (Figs. 9, 13 C) Radula
11. Dorsal lobe with 2 diverging lobules (Fig. 12) Goebeliella
Ventral lobule and segments of stipules often saccate Lepidolaena
12. Leaf margins lobed, sometimes toothed, stipules large, resembling leaves Isotachis
Leaves transverse, concave-complicate, shortly bi-lobed 13
Leaves incubous with long tapering lobes 14
Leaves shaped otherwise 15
13. Leaves imbricate, sometimes papillose, perianth ventral, basal Lembidium (excluding two dendroid species)
Plants weak, hyaline, perianth terminal Hygrobiella
14. Leaves bi-lobed, homomallous Herberta
Leaves twice bi-lobed or 3-5-lobed, dorsal margin sometimes dentate-ciliate Lepicolea
15. Leaves unevenly 2-lobed, plant pendulous, flagelliferous Mastigophora
Leaves 3-6 deeply cleft 16
Leaves usually flat, longer than mid-width, apex 2-3-toothed 18
16. Perianth scaly, leaves with numerous hair-like divisions Trichocolea
Perianth smooth 17
17. Branches sometimes attenuated, leaf-segments usually entire, sometimes toothed (Fig. 8) Lepidozia
Branches not attenuated, leaf-segments fringed with long cilia Ptilidium
18. Leaf-apex usually 3-toothed, ventral branches from the axils of the stipules (Figs. 4, 5) Bazzania
Leaf-apex 2-dentate or 2-lobed (entire in an undescribed species), ventral branches from the side of the stipules Acromastigum
19. Leaves with a smaller dorsal lobe (equal in Diplophyllum densifolium) 20
Leaves without a dorsal lobe 22
20. Stipules absent (Fig. 2) Diplophyllum
Stipules present 21
21. Dorsal lobe usually broader than tall, sporophyte in a terminal fleshy cup Schistochila
Dorsal lobe usually taller than, or as tall as broad, sporophyte terminal in a marsupium (Fig. 6) Balantiopsis
22. Cauline stipules present 23
Cauline stipules absent (except in Jungermannia rotata) 29
23. Leaves 3-6 deeply cleft 24
Leaves shaped otherwise 25
24. Leaf-segments spiny, in one plane, perianth terminal Blepharostoma
Leaf-segments erect from a spreading base, perianth lateral Psiloclada
25. Perianth terminal on main stem (rarely lateral in Lophocolea) 26
Perianth or marsupium lateral (rarely terminal in Chiloscyphus) 27
26. Perianth 3-gonous, third angle dorsal (Fig. 10) Lophocolea
Perianth laterally compressed, mouth 2-lipped Mylia
Perianth deeply multiplicate, leaves undulate, bilobed, plants robust Chandonanthus
27. Sporophyte in a campanulate perianth (Fig. 7) Chiloscyphus
Sporophyte in a fleshy marsupium 28
28. Leaves and stipules deeply bifid Geocalyx
Leaf-apices entire or irregularly sub-entire, papillose Saccogyna
29. Sporophyte not in a marsupium, usually a perianth 30
Sporophyte in a marsupium 40
30. Perianth terminal on main stem or branches 31
Perianth remote from apex (sometimes terminal in Cephalozia), branches ventral 39
31. Perianth laterally compressed, leaves usually toothed (Fig. 30, O) Plagiochila
Perianth shaped otherwise or absent (in Gymnomitrium) 32
32. Leaves bilobed 33
Leaves entire 37
33. Plants robust, reddish Anastrophyllum
Plants minute to small 34
34. Perianth absent, leaves closely imbricate, complicate, only shortly bilobed Gymnomitrium
Leaves sub-remote to remote 35

Legends to Figures Plate 1 1. Diplasiolejeunea lyratifolia ventral, showing duplicated stipules, one for each leaf, × 10. 2. Diplcphyllum domesticum dorsal, showing small dorsal leaf-lobes, × 5. 3. Strepsilejeunea Curnowii ventral, showing acute, decurved leaf-apices, × 10. 4. Bazzania Novae-Zelandiae ventral, showing incubous leaves and ventral axillary branching, × 4. 5. Bazzania Novae-Zelandiae dorsal, showing incubous leaves, × 4. 6. Balantiopsis diplophyllum dorsal, showing terminal marsupium, × 6. 7. Chiloscyphus ammophilus dorsal, showing succubous leaves, × 5. 8. Lepidozia tetradactyla dorsal, showing deeply divided leaves and lateral perianth, × 8. 9. Radula Levieri ventral, showing lobules, absence of stipules and spent capsule, × 10. 10. Lophocolea subporosa dorsal, showing terminal perinath with 3rd keel dorsal, × 5. 11. Cephaloziela sp., dorsal, showing deeply pluri-plicate perianth and bi-fid leaves, × 12. 12. Goebeliella cornigera ventral, showing double lobules and entire stiples, × 10 13. Elaters, A Metzgeria violacea monospiral, × 125; B. Frullania rostallata monospiral; C. Radula Levieri bispiral. 14. Pseudolates of Anthoceros × 100. 15. Cells of Lejeunea sp. without trigones, ca. 40 microns. 16. Rhizoids; D. Reboulia hemispherica, smooth; E. Lunularia cruciata, tuberculate, × 125.

Legends to Figures
Plate 1
1. Diplasiolejeunea lyratifolia ventral, showing duplicated stipules, one for each leaf, × 10.
2. Diplcphyllum domesticum dorsal, showing small dorsal leaf-lobes, × 5.
3. Strepsilejeunea Curnowii ventral, showing acute, decurved leaf-apices, × 10.
4. Bazzania Novae-Zelandiae ventral, showing incubous leaves and ventral axillary branching, × 4.
5. Bazzania Novae-Zelandiae dorsal, showing incubous leaves, × 4.
6. Balantiopsis diplophyllum dorsal, showing terminal marsupium, × 6.
7. Chiloscyphus ammophilus dorsal, showing succubous leaves, × 5.
8. Lepidozia tetradactyla dorsal, showing deeply divided leaves and lateral perianth, × 8.
9. Radula Levieri ventral, showing lobules, absence of stipules and spent capsule, × 10.
10. Lophocolea subporosa dorsal, showing terminal perinath with 3rd keel dorsal, × 5.
11. Cephaloziela sp., dorsal, showing deeply pluri-plicate perianth and bi-fid leaves, × 12.
12. Goebeliella cornigera ventral, showing double lobules and entire stiples, × 10
13. Elaters, A Metzgeria violacea monospiral, × 125; B. Frullania rostallata monospiral; C. Radula Levieri bispiral.
14. Pseudolates of Anthoceros × 100.
15. Cells of Lejeunea sp. without trigones, ca. 40 microns.
16. Rhizoids; D. Reboulia hemispherica, smooth; E. Lunularia cruciata, tuberculate, × 125.

35. Perianth pluriplicate, branches ventral (Fig. 11) Cephaloziella
Perianth ovate, mouth plicate, branches lateral 36
36. Leaves spreading Sphenolobus
Leaves (more or less) concave Lophozia
37. Leaf-apex acute or cuspidate Cuspidatula
Leaf-apex rounded 38
38. Medium to robust, leaves erect, imbricate, invol. stipule present Jumesoniella
Small to medium, leaves soft and crumpled, invol. stipule usually absent Jungermannia
39. Plants small, perianth trigonous, third angle ventral, leaves two or more lobed Cephalozia
Plants medium to sub-robust, perianth cylindrical, leaves not lobed Adelanthus
40. Marsupium lateral Marsupidium
Marsupium terminal 41
41. Stems erect, marsupium pendulous, hairy Tylimanthus
Stems prostrate, marsupium smooth 42
42. Leavse entire Symphyomitra
Leaves bilobed Acrobolbus
43. Sporophyte sunk in the tissues of the thallus 44
Sporophyte not sunk in the tissue of the thallus 45
44. Epidermis without pores, antheridia scattered Riccia
Epidermis without pores, antheridia in a median furrow Ricciocarpus
45. Sporophyte an erect linear organ of two valves dehiscing downwards 46
Archegonia in other combinations from the thallus 49

17. Asterella australis × 16, F carpocephalum showing laciniate perianth, × 4. 18. Targionia hypophylla with terminal valves, × 1½. 19. Riccardia marginata showing absence of nerve and lateral calyptra, × 3. 20. Anthocercs with linear dehiscing capsule, nat. size. 21. Marchantia with male carpocephalum and rounded gammae cups; G. female carpocephalum showing involucres between the rays, nat. size. 22. Metzgeria furcata dorsal with ventral calyptras showing, also mid-rib and ciliate margins; H. ventral with male branches, x. 4. 23. Reboulia hemisphaerica, carpocephalum, underneath view showing bi-valved involucres, × 2½. 24. Plagiochosma australe with invol. lobes opening vertically, × 2½. 26. Lunularia cruciata with lunate gemmae cups, nat. size. 27. Sypmhyogyna hymenophyllum showing involucral scales and calyptras; I. sterile archegonia adhering to calyptra, × 2. 28. Pallavicinia Lyellii, J. invol. cup, K. perianth with calyptra included, nat. size. 29. Hymenophytum phyllanthus, L. basal ventral invol. cup, M. perianth, calyptra included, nat. size. 30. Dehisecnt capsules; N. Taxilejeunea Colensoana showing valves not divided to the base, × 10; O. Plagiochila × 7; P. Fossombronia × 7; Q. Metzgeria furcata with elaterophones × 10; R. Asterella × 5.

17. Asterella australis × 16, F carpocephalum showing laciniate perianth, × 4.
18. Targionia hypophylla with terminal valves, × 1½.
19. Riccardia marginata showing absence of nerve and lateral calyptra, × 3.
20. Anthocercs with linear dehiscing capsule, nat. size.
21. Marchantia with male carpocephalum and rounded gammae cups; G. female carpocephalum showing involucres between the rays, nat. size.
22. Metzgeria furcata dorsal with ventral calyptras showing, also mid-rib and ciliate margins; H. ventral with male branches, x. 4.
23. Reboulia hemisphaerica, carpocephalum, underneath view showing bi-valved involucres, × 2½.
24. Plagiochosma australe with invol. lobes opening vertically, × 2½.
26. Lunularia cruciata with lunate gemmae cups, nat. size.
27. Sypmhyogyna hymenophyllum showing involucral scales and calyptras; I. sterile archegonia adhering to calyptra, × 2.
28. Pallavicinia Lyellii, J. invol. cup, K. perianth with calyptra included, nat. size.
29. Hymenophytum phyllanthus, L. basal ventral invol. cup, M. perianth, calyptra included, nat. size.
30. Dehisecnt capsules; N. Taxilejeunea Colensoana showing valves not divided to the base, × 10; O. Plagiochila × 7; P. Fossombronia × 7; Q. Metzgeria furcata with elaterophones × 10; R. Asterella × 5.

46. Thallus of one cell in thickness, with a thicker vein Dendroceros
Thallus several cells thick without a vein 47
47. True spiral elaters present in the capsule Megaceros
Pseudoelaters present in the capsule 48
48. Capsule walls with stomata (Figs. 14, 20) Anthoceros
Capsule walls without stomata Aspiromitus
49. Thallus divided into layers of different tissue with pores from air-cavities 50
Thallus not divided into layers, and without pores and air-cavities 55
50. Archegonial group terminal, enclosed in two scales (Fig. 18) Targionia
Archegonia on peduncled receptacles (carpocephala) 51
51. Involucre with one capsule, dehiscing by the falling of a lid, leaving a cup 52
Involucre with a group of capsules (one, with infertile archegonia in Lunularia) dehiscing by 4-8 valves 54
52. Carpocephalum dorsal, involucres ascending with a vertical aperture, ventral scales large, hyaline-tipped, upper projecting beyond the margin (Fig. 24) Plagiochasma
Carpocephalum terminal, ventral scales not hyaline-tipped 53
53. Involucres 2-valved, no perianth (Figs. 16, D, 23) Reboulia
Involucres not 2-valved, a white basket-like perianth present, split into fimbriae (Figs. 17, 30 R) Asterella
54. Peduncle not grooved, gemmae cups lunate, fruiting plants very rare (Fig. 16 E, 26) Lunularia
Peduncle grooved, gemmae cups round when present (Fig. 21) Marchantia
55. Thallus sometimes gigantic, densely rhiziferous, archegonia terminal, capsule 1-valved splitting on one side and opening till flat (Fig. 25) Monoclea
Archegonia not terminal, capsule dehiscing by 4 valves ( Jung. anacrogynae) 56
56. Sexual organs from the upper surface of the frond 57
Sexual organs not from the upper surface 62
57. Frond lamellate 58
Sterile and female frond not lamellate 59
58. Frond roundish, perianth with a wide mouth Petalophyllum
Frond elongate, sectioned off into succubous lobes, with a lamella at the dorsal base of each, perianth none Treubia
59. Involucre of laciniate scales, perianth present 60
Involucre of laciniate scales, perianth absent 61
60. Stipules and elaterophores absent (Fig. 28) Pallavicinia
Stipules and elaterophores present Calycularia
61. Invol. scales at the lower base of the calyptra only, calyptra fleshy, no elaterophores (Fig. 27) Symphyogyna
Scales forming a (more or less) complete ring, large and much lacerated, calyptra shortish, sub-globose, elaterophores present Allisonia
62. Sporophyte ventral, but just below the membraneous expansion, or at the lower fork, involucre and perianth present (Fig. 29) Hymenophytum
Sporophyte not basal, perianth absent 63
63. Sexual organs on short ventral branches, frond margins often ciliate, cells unistratose except at midrib (Figs. 13A, 22, 30 Q) Metzgeria
Sexual organs on short marginal branches, not ciliate, cells usually multistratose, no mid-rib (Fig. 19) Riccardia

Key To New Zealand Genera

* In Descriptions of New Species of N.Z. Hepatics, Trans. Roy. Soc. N.Z., 1938, pp. 45-46.

1. Stipules present 2
Stipules absent 19
2. Stipules duplicated, one for every leaf 3
One stipule for every two leaves 4
3. Dorsal lobe bigger than the ventral, not papillose Diplasiolejeunea
Ventral lobe bigger than the dorsal—a tubular lobule, widening out into a large sack, strongly papillose Colura
4. Stipules entire, not bifid nor two-lobed 5
Stipules bifid or two-lobed 9
5. Lobules and apex of leaves and stipules toothed (rare) Ptychanthus
Leaves and stipules not toothed 6
6. Perianth 3-gonous, leaves densely imbricated, stems noticeably constricted at intervals, stipules narrowed to the base Thysananthus
Perianth not 3-gonous, leaves not usualy densely imbricated, stipules rounded to the base 7
7. Plants large, usually dark green drying to dingy brown, leaves fairly round (moist), perianth not immersed, oblong, dorsally flat with ribs Archilejeunea
Plants smaller, perianth immersed or slightly emergent 8
8. Leaves drying to glossy brown, sub-falcate, perianth dorsally flat and round, margins toothed or leafy, with innovations Lophololejeunea
Leaves yellow-brown, squarrose when moist, perianth globose, deeply 10-keeled with no innovations Ptychocoleus
9. Plants filiform, white, leaves scarcely contiguous, to remote 10
Plants small to medium, leaves usually imbricate or subimbricate 12
10. Leaves obtuse, stipules round Microlejeunea
Leaves acute to acuminate, triangular in general outline, erecto-patent, stipules broader at the apex of the segments 11
11. Stipule segments obtuse, sinus shallow Harpalejeunea
Sipule segments acute, widely diverging Drepanolejeunea
12. Leaves with high papillae, whitish, usually on filmy ferns Trachylejeunea
Leaves not, or lowly papillose 13
13. Leaves and lobules 2-3 times longer than broad 14
Leaves rarely more than 1½ times longer than broad 15
14. Leaves narrowly obovate or sub-lanceolate, margins crenulate, perianth with a long neck (common) Siphonolejeunea
Leaves squarrose, narrow-oblong, concave, cells large, irregular, thick-walled (rare) Leptolejeunea
15. Ventral margin of the lobe with no indentation at the junction with the lobule, enlarged cells (ocelli) present, perianth dorsally compressed Rectolejeunea
Lobule not in a straight line with the ventral margin of the lobe, (except perhaps in Lejeunea), perianth not dorsally compressed 16
16. Leaf-apex usually acute, decurved (Fig. 3) Strepsilejeunea
Leaf-apex rounded 17
17. Lobule small, cells transparent without trigones (Fig. 15) Lejeunea
Lobule medium to large, trigones present 18
18. Perianth smooth, inflated, with repeated innovations (Fig. 30N) Taxilejeunea
Perianth 5-keeled, lobule involute (more or less) cylindric, plants paler Pycnolejeunea
19. Minute, lobule large (comparatively), sub-globose, perianth 5-plicate, papillose Cololejeunea
Small, whitish, lobule flat, 2-3-dentate, perianth compressed, obcordate, plants not papillose Leptocolea

A Glossary of Terms Used in This Paper.