Lepidozia pendulina (Hook.) Lindenb.
Jungermannia pendulina Hook., Musci Exot. 1: pl. 60, f. 1–5. 1818.
Lepidozia pendulina (Hook.) Lindenb. in Gottsche, Lindenb. & Nees, Syn. Hepat. 208. 1845.
Mastigophora pendulina (Hook.) Trevis., Mem. Reale Ist. Lombardo Sci. Lett. III, 4: 416. 1877.
Type: New Zealand, South Is., Dusky Bay, 1791, Menzies, ex herb. Kew (G!).
Lepidozia gigantea Steph., Sp. Hepat. 3: 600. 1909.
Type: New Zealand, Dall “Doll” (G!).
[Plate 5D; Fig. 34: 2, oil-bodies, p. 220]
Plants erect, rigid, with ventrally secund branches, pale green, the shoots robust, to 6 cm wide, including branches. Branching mostly of Frullania type, consistently 2-pinnate (rarely 3-pinnate), the branches whip-like, often becoming flagelliform, the leaves like those of main shoot except smaller; branch half-leaf 2-lobed or sporadically unlobed; first branch underleaf (3)4-lobed, inserted on ventral-lateral side of main axis in a plane similar to leaves of main shoot. Ventral-intercalary branching rare and sporadic. Stems rigid, the cortical cells in 1 layer of firm-walled cells, internal to the epidermal layer are 2–3 to 4–6 rows of thick-walled cells; central portion of stem made up of cells larger in diameter and with thin walls. Leaves rigid, slightly concave, distant, 1.1–1.4 mm long at longest point, (1.7)1.9–2.5 mm wide at widest point, stiffly patent, the insertion transverse; leaves asymmetric, unequally 4(6)-lobed, the leaves divided to ca. 0.65–0.75 (median sinus), the distance from dorsal sinus base to insertion much greater than that from ventral sinus to insertion. Lobes broadly attenuate (the dorsal lobes acute, the ventral lobe more narrowly attenuate and curved), the dorsal pair of lobes partially united, the others distinct and spreading, the ventral lobe often widely divergent as a “claw,” the dorsal lobes entire, (10)14–24 cells wide at base, the lobes terminating in a uniseriate row of 2–4(6) cells (6–7 in ventral-most lobe); cells of uniseriate row ± isodiametric to slightly longer than wide, thick-walled. Disc distinctly asymmetric, 20–30 cells high at dorsal sinus, 12–18 cells high at ventral sinus, the margins entire, the dorsal margin broadly ampliate, the ventral slightly curved. Cells of disc-middle thick-walled, trigones medium and straight-sided, occasionally with intermediate thickenings, ± isodiametric to elongate, 18–26(29) µm wide × 24–33(42) µm long, in ± regular longitudinal files; median basal cells enlarged, in 1–2 rows, with walls often nodulose; surface smooth to indistinctly striate. Oil-bodies occupying a small portion of cell lumen, hyaline, (2)3–4(5) per cell at base of dorsal sinus (in median cells (2)3–8 per median cell, locally to 10–14 per cell, fide Engel and Schuster, 2001), globose to elliptic to fusiform, a few linear in profile, botryoidal, but quickly breaking up by becoming increasingly botryoidal via swelling of spherules, 4.3–4.8 × 7.7–8.6 µm to 4.3–6.2 × 10.6–12.5 µm, globose ones 4.3–4.8 µm in diam. Underleaves often weakly to distinctly plicate, widely spreading, symmetrically 4(6)-fid to ca. 0.65–0.7 (median sinus), the sinus bases somewhat reflexed, the lobes ± parallel, somewhat ventrally sulcate, narrowly and gradually attenuate, entire, terminating in a uniseriate row of 1–3(4) cells; disc margins reflexed, entire or sporadically with a tooth. Fungal partner an ascomycete.
Plants dioecious. Androecia on inconspicuous, short, determinate, tightly spicate, cernuous ventral-intercalary branches from main shoot and primary + secondary branches (in flagelliform or leafy sectors); bracts ventricose-cucullate, 2-lobed to ca. 0.3, the lobes acute to short-acuminate; antheridial stalk biseriate. Gynoecia only sporadically produced, on abbreviated ventral-intercalary branches issuing from main stem; bracts of innermost series deeply concave, broadly ovate-subrectangular; apices with 4–5 short, ± regular lobes, the lobes finely crenulate; lamina margin bordered by thin-walled cells of variable shape and orientation, the cells often ± sinuate-rhomboidal, the apical or free end of marginal cells variously divergent and forming a short projection or a tooth, the margin irregularly crenulate to crenate-denticulate; bracteole similar in size and form. Perianth long and prominent, slenderly cylindrical-fusiform, slightly curved, terete below, obscurely trigonous above, distinctly and deeply 3-plicate toward mouth, the perianth gradually narrowing toward the strongly contracted, shallowly 3-lobed mouth, the lobe margins composed of irregularly sinuate-rhomboidal cells that, at the apical end, project for varying lengths, the mouth thus crenate-denticulate.
Seta with 16 rows of outer cells surrounding an inner core of numerous much smaller cells. Mature capsules not seen.
Distribution and Ecology : New Zealand: Stewart Island (5 m), South Island (0–1200 m), North Island (1000–1200 m); Australia: Tasmania, Victoria. Occurs in mainly western, high rainfall parts of the country but extends to the centers of both main islands. Found in Fiordland, Southland, Westland, western edges of Canterbury, Western Nelson, Southern North Island (Tararua and Ruahine ranges), Taranaki (Egmont Natl. Park), Volcanic Plateau (Tongariro Natl. Park) and Gisborne (Urewera Natl. Park) EPs. Apparently absent from Auckland and Northland provinces.
The species occurs in dense, very wet to only moderately wet forests over a broad altitudinal gradient, but is more typical of middle- to upper-elevation sites. It occurs in Nothofagus solandri, N. menziesii and N. truncata forest. Other species such as Metrosideros umbellata and Weinmannia racemosa may also be present as canopy species, but Lepidozia pendulina seems to be confined to forests in which Nothofagus is present. For example, it may occur at the upper limits of N. menziesii forests (stations at ca. 900 m in the South Island, and ca. 1180 m in the North Island) as well as in upper montane N. solandri forests with Phyllocladus alpinus, Brachyglottis buchananii, Pseudopanax spp., Dracophyllum traversii and Coprosma. In the southern sectors of South Island it occurs in forests at lower elevations including at sea level.
The species typically occurs over organic substrates such as the forest floor (particularly in damp leaf litter) where it may form extensive, at times large, pure carpets that at times raise to low-lying mounds; it may be associated with Bazzania nitida, Lepidogyna hodgsoniae, Plagiochila gigantea, Schistochila nobilis and Trichocolea mollissima. It also occurs on damp, bryophyte-covered banks and wet, bryophyte-covered, rotted logs. The species often is a component of the flora of sites where the floor, boulders and trees are densely covered with bryophytes. Associated species are: Adelanthus falcatus, Bazzania adnexa, Blepharidophyllum vertebrale, Breutelia pendula, Chiloscyphus mittenianus, Cryptochila grandiflora, Dendroligotrichum dendroides, Dicranoloma billardierei, D. robustum, Gackstroemia weindorferi, Herbertus oldfieldianus, Heteroscyphus cymbaliferus, H. sinuosus, Hymenophyllum multifidum, Lepidogyna hodgsoniae, Leptotheca gaudichaudii, Paraschistochila pinnatifolia, Plagiochila gigantea, P. retrospectans, Pogonatum subulatum, Pseudocyphellaria multifida, Ptychomnion aciculare, Pyrrhobryum bifarium, Rhizogonium distichum, Schistochila appendiculata, S. ciliata, S. glaucescens, S. nobilis, Trichocolea mollissima, Tylimanthus saccatus and Wijkia extenuata.
Comments : Well-developed plants of this species are not likely to be confused with any other New Zealand member of the genus on general aspect alone. Distinctive are the bipinnate branching (small, poorly developed populations have secondary branches weakly developed or absent); the broad, transversely inserted, entire-margined leaves; the large underleaves with ± parallel, ventrally sulcate lobes; and the leaf areolation, the leaf cells thick-walled, arranged in ± regular longitudinal files, and appearing small and narrow in proportion to the size of the leaf, giving the leaf a densely cellular appearance, even under the dissecting microscope.
