Herbertus oldfieldianus (Steph.) Rodway
Type: Australia, Tasmania, Oldfield s.n. (G!); isotype: (FH!).
Schisma alpinum Steph., Sp. Hepat. 4: 20. 1909.
Herberta alpina (Steph.) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3: 185. 1967.
Type: New Zealand, W side of Ruahine Ra. Colenso 5179 (G!, WELT 7619!).
[Plate 2D; Figs. 11, 12; Fig. 13: 1, 2, oil-bodies, p. 114]
Plants often forming compact tufts, stiff, olive-green to yellow-brown, at times tinged with red-brown, with a system of branched, geotropic and plagiotropic stolons and microphyllous axes from which arise erect, leafy shoots that are often feebly cernuous at the tips; shoots 20–40 mm long, to 2 mm wide, the older parts of the shoots often bare of leaves. Branches of leafy shoots common, ventral-intercalary, often stoloniform, often initially small-leaved and soon becoming flagelliform. Leaves rigid, ventrally secund, imbricate, the insertions overlapping dorsally, the dorsal bases of opposing leaves distinctly overlapping and covering stem in dorsal view, divided to 0.65–0.75, 0.50–0.8 mm wide at the lobe bases, 0.6–1.05 mm wide between the lobe apices, 1.3–1.8 mm long; lobes falcate, 2.9–4.8× longer than wide, convex toward base (sometimes strongly so on the dorsal side), the distal sector (especially of ventral lobe) concave-channeled, the tip often apiculate, terminating in a single cell or a uniseriate row of 2–5 cells; lamina convex, the margins inflexed, with several sessile slime papillae; vitta of leaves distinct, extending nearly to lobe apices. Leaf cells with trigones massive, the marginal and submarginal cells of lamina 17–22 µm wide, 11–18 µm long, the intramarginal cells becoming gradually larger as the vitta is approached; vitta cells near leaf base with transverse radial walls very thin, the cells 22–25 µm wide, 50–70 µm long; lobe tip surfaces rather distinctly striate-papillose, of disc obscurely so. Oil-bodies appearing rather crowded in vitta cells, in non-vitta cells occupying only moderate portion of cell, pale smokey grey or hyaline and glistening, 13–15 per vitta cell, those in vitta cells globose to narrowly to broadly elliptic, occasionally sublinear, 4 × 4 µm to 3 × 7 µm; oil-bodies in nonvittate cells 3–6 per cell, 1.5–3 × 2.5–4 µm; oil-bodies breaking up by way of tiny spherules that swell from the oil-body surface. Underleaves similar in size and form to leaves but not secund, widely spreading to squarrose, oblong-elliptic to cuneate; lobes ± parallel to somewhat diverging; lamina margins abruptly incurved, the several slime papillae sessile or with a unicellular stalk. Asexual reproduction lacking or ?by fragmenting leaves and ♀ bracts. Fungal partner absent.
Androecia on main shoot, wider than sterile sectors of shoot; bracts more closely imbricate than leaves, in 4–6 pairs, the lobes falcate, the bracts and bracteoles less deeply lobed than leaves, the lamina distinctly ventricose, the lamina margins irregularly denticulate, at times sparingly so, the margins with sessile slime papillae. Gynoecia sporadic, on main shoot; bracts and bracteoles similar (except bract lobes feebly falcate), forming a compact and “bud-like” structure, those of innermost series with at least the basal sector appressed to and ensheathing perianth, divided to ca. 0.5 by caudate lobes, the lamina deeply concave, the shape of the concavity(ies) conforming to the shape of perianth plicae, margins of lamina and lower sectors of lobes with spinose teeth and copious sessile and stalked slime papillae, the stalks + spinose teeth rendering the margins irregularly denticulate-dentate. Perianth barely exserted beyond bracts, 3.2–4.1 mm long, 1.2–1.3 mm in diam., ± elliptic, basically trigonous but each angle with 2 lobes, divided to the middle into 6 caudate lobes ca. 1.5 mm long, the lobe margins denticulate-dentate and with copious slime papillae.
Seta ca. 3.2 mm long. Capsule splitting longitudinally but unevenly, the wall 6–8-layered and 48–60 µm thick to 8–9-layered and 86–90(100) µm thick (Schuster, 2000a), the outer layer of cells equal to thickness of slightly over 2 of interior strata; outer layer of cells densely covered with an amorphous, ± granular, waxy substance (soluble in xylene), irregular in shape, subquadrate to irregularly short-rectangular, the radial walls firm, hyaline, with red-brown, strongly nodular thickenings on both longitudinal and transverse walls, the thickenings often appearing “stalked,” the nodules rather irregular in shape, some with a notch at the summit, some attenuated and spine-like, most pigmented, a few hyaline, a few band-like thickenings present at the cell ends or edges; innermost layer of cells irregularly narrowly elongate-rectangular, the radial walls with nodular thickenings common, occasionally with semiannular bands.
Spores 19.2–20.6 µm in diam., reddish brown, tuberculate-baculate, the tubercles truncate and often dilated at the summit, the ground surface between tubercles appearing smooth. Elaters in number adhering by one end to inner face of capsule, feebly tortuous to nearly straight, 9.6–10.6 µm wide, 3–4-spiral in median portion, bispiral and considerably more loosely wound toward ends, the spirals 2.4–2.9 µm wide; elaters at times bispiral throughout.
Distribution and Ecology : New Zealand: Campbell Island, Stewart Island, South Island (440–1540 m), North Island (960–1330 m); Australia: Tasmania. In New Zealand known from Fiordland, Otago (Mt. Cargill, Garvie Mtns.), Canterbury (Arthur’s Pass, Governors Bush), Westland, Western Nelson, Sounds–Nelson (Mt. Stokes, Mt. Richmond), Southern North Island (Ruahine Ra., Tararua Ra.), Volcanic Plateau, Taranaki (Mt. Taranaki), Gisborne and Auckland (Coromandel Peninsula) EPs. The northern limit is at Mt. Moehau on Coromandel Peninsula. In the South Island, common west of and on the Main Divide but not extending far to the east.
A plant chiefly of upper montane forest and scrub of Nothofagus menziesii, N. solandri var. cliffortioides, Archeria traversii, Dracophyllum traversii, D. longifolium, D. rosmarinifolium, and Olearia colensoi where it is usually epiphytic on hosts that include N. menziesii, N. solandri, Halocarpus bidwillii and Weinmannia racemosa, and is particularly common on Archeria traversii. It is also found on soil and rock under forest and scrub. It forms large, compact tufts on tree bases and may be locally abundant and crowded in such situations and form polsters. It also occurs in open sites on ledges and crevices of rocky (often granite) outcrops, boulder faces and steep slopes in the penalpine and alpine zones. It also occurs on damp, shady banks and on rocky knolls of slopes of mostly exposed, bare rock in areas with sparse vegetation (e.g., consisting of a few scattered shrubs and grasses). Herbertus oldfieldianus may occur near sea level (e.g., at Milford Sound). It is often found with Acrobolbus cinerascens, Anastrophyllum schismoides, Dicranoloma billardierei, Goebeliella cornigera, Jamesoniella colorata, Leifidium tenerum, Lepicolea attenuata, L. scolopendra, Lepyrodon australis, Paraschistochila pinnatifolia, P. tuloides, Plagiochila circinalis, Plagiochilion conjugatum and Radula pseudoscripta, and, when on rock, with Acrolophozia pectinata, Andreaea acutifolia, Cheilolejeunea campbelliensis, Frullania aterrima, Jamesoniella colorata and Rhacocarpus purpurascens.
Comments : Large, fuscous or olive-brown phases typically form tufts or cushions, often extensive and pure; such plants may be many centimeters tall and have secund shoot apices, much as in the moss Dicranoloma. With the three rather dense series of deeply bifid leaves, the shoots have a distinctly moss-like aspect—the long-tapered leaf lobes simulating leaves of a moss. Such plants copiously produce slender, often whip-like, frequently ramified, chiefly geotropic flagellae that simulate a root system.
By contrast, alpine/penalpine phases may be much more compact, only 1–2 cm tall × 1.2–1.5 mm wide with the leaves; such phases may be reddish brown in exposed sites, dense-leaved, with lateral leaves only weakly falcate and with less acuminate lobes. These plants have scarcely secund shoot apices and may also closely mimic mosses.
Similar to Triandrophyllum, especially the very stunted forms found on exposed rock. The leaves are more imbricate and less spreading from the stem, the stems are wiry and often bare of leaves at their base. Also similar to Lepicolea and Dendromastigophora, but the branching is not pinnate and branches are at a very narrow angle rather than at right angles to the stem as they are in those two genera. The cushion-forming habit of the species, whether epiphytic or terrestrial, is distinctive, as are the secund leaves, a common feature of mosses, but rarely seen in liverworts.
