Trichocolea rigida R.M.Schust.
Trichocolea rigida R.M.Schust., Nova Hedwigia 15: 447. f. 50: 1–7. 1968.
Holotype: New Zealand, South Is., Otago Prov., Old Man Ra., ca. 4000 ft., Schuster 67-523 (herb. Schuster); isotypes: (herb. Grolle, NICH, OTA, S).
[Fig. 24: 3, oil-bodies, p. 178; Fig. 25]
Plants relatively rigid, woolly in appearance, prostrate, densely interwoven, yellow-green, whitish with age, the dried plants dull, appearing amorphous and resembling a crustose lichen, the shoots small, to 1 cm wide, including branches. Branching irregularly 2-pinnate, with 2–4(5) ± irregularly curved, secondary branches per primary branch; branches julaceous, with a woolly appearance, with all sides appearing as a mass of interwoven cilia. Stems rather stiff, paraphyllia rather abundant, much branched, with cells similar to those of the leaf cilia. Rhizoids usually absent. Leaves of stem widely spreading but with lobes and cilia broadly arching upward toward shoot apex, the leaf thus somewhat hand-like, the leaves in situ appearing as a dense mass of cilia, the individual lobes hardly discernable, the leaves on the stem approximate, becoming imbricate on branches, semiamplexicaul, the insertion transverse to weakly succubous, obtrapezoidal to subreniform, divided into 4–5 erect, pinnulate lobes, the lobes sometimes appearing forked or partially fused at base (those in ventral sector of leaf somewhat larger), each lobe actually unequally and irregularly (2)3-fid, consisting of a stronger, erect, pinnulate division and (0)1–2 small, freely branched and variably positioned accessory divisions that are 1–2(3) cells wide at the base, and stiffly abaxially spreading and arched in same direction as the main lobes, the divisions armed with pairs of opposing branched cilia. Lobes caudate, ± regularly pinnulate (best seen in adaxial view), with numerous stout segments and cilia, the ventral lobes 4–6 cells wide at base, with 3–5 tiers of laterally juxtaposed, turgid cells basal to the uniseriate row of (4)5–7(8) cells; cells of uniseriate row short, barrel-shaped, rather thin-walled and ± turgid, constricted at the septa, the septa not thickened or dilated, the surface coarsely papillose, the penultimate cell (14)17–24 µm wide, (43)50–67(74) µm long; terminal cell of uniseriate row finger-like, slightly tapering to a broadly rounded apex, 12–22 µm wide, 50–65(82) µm long, thin-walled in the tip, the surface distinctly papillose; margins of lobes armed with pairs of abaxially displaced, opposing cilia, the cilia usually branched (often repeatedly so), the armature all inserted on the abaxial surface of lobe and abaxially patent, but ascending and nearly as long as lobes; marginal cilia of lobe mostly uniseriate throughout, the cilia bases rarely with 2 laterally juxtaposed cells, the cells similar to those of the uniseriate sector of leaf lobes; sinus bases plane and not reflexed. Disc weakly asymmetric, consisting of a low, wide strip 3–4 cells high (from base to sinuses); margins of disc with freely branched cilia, the cilia cells similar to those of lobes, the cilia sometimes abaxially displaced and lying at right angles to the disc plane. Cells of disc faintly uniformly thick-walled, in lamina middle 22–34 µm wide, 53–75(80) µm long; surface densely striate-papillose to striolate. Oil-bodies within all cells of leaf including tip cell of cilia, occupying small portion of cell lumen, opaque, smokey grey, 4–8(12) per paired cells at lobe base, irregularly and inconspicuously and finely granular, the spherules tiny and irregular, the oil-bodies irregular in shape: globose to bottle-shaped to elliptic to sublinear to subcrescentic to subsigmoid, 4.3–4.8 × 7.2–8.6 µm to 2.9–4.8 × 9.6–11.5 µm or spherical and 4.8 µm in diam. Underleaves in situ appearing as a dense mass of cilia, somewhat smaller than leaves, free, suborbicular to reniform, quadrifid, often weakly bisbifid, the armature similar to leaves; disc 2–3 cells high.
Androecia intercalary on primary and secondary branches, appearing brushed with apically oriented cilia, the bracts divided to ca. 0.4–0.5, the lamina strongly ventricose, the abaxial face of bract with simple or sporadically branched cilia in distal sector, the armature only sporadically on lamina; antheridia large for bract size, 1(2) per bract, the stalk biseriate. Gynoecia and sporophyte not seen.
Distribution and Ecology : New Zealand: Stewart Island (10–30 m), South Island (0–1390 m), North Island (200–1250 m), Chatham Islands (240 m); Australia: Tasmania and Victoria. In New Zealand known from Northland, Auckland, Volcanic Plateau, Southern North Island (Ruahine and Tararua ranges), Western Nelson (Cobb Valley), Westland, Canterbury (Banks Peninsula, Hawdon River), Otago (Central Otago, Lammerlaw and Otago Coast ERs), Fiordland and Rakiura (Stewart Island, Solander Islands) EPs.
In New Zealand the species is known from scattered localities over a broad altitudinal range. It occurs on the floor and mossy banks in forests near the sea (Martin’s Bay and Milford Sound in Fiordland), as well as in upper-elevation forests (e.g., Nothofagus fusca – N. solandri var. cliffortioides forest at ca. 1280–1390 m at Pinchgut Track, Nelson Lakes Natl. Park). It also occurs in the penalpine zone in damp or springy niches under snow tussock (e.g., in the Lammermoor Ra., Old Man Ra., the type, both in Otago). Also on roadside banks at Halfmoon Bay, Stewart Island. Unlike our two other species, which occur in forests, Trichocolea rigida commonly occurs in more open sites, e.g., in seepage areas in penalpine tussock grassland (Chionochloa) and exposed roadside banks, and in grassland kept damp by seepage, for instance at Gebbies Pass and Hilltop on Banks Peninsula. At Stockton it occurs under gorse scrub in a gully, on damp soil with adventive grasses and Polytrichum commune. On Chatham Island it was found on a soil bank under a canopy of Dicksonia fibrosa. Species found with T. rigida are Brachythecium rutabulum, Breutelia pendula, Chiloscyphus leucophyllus, Cratoneuropsis relaxa, Distichophyllum pulchellum, Drepanocladus aduncus, Fissidens asplenioides, Canalohypopterygium tamariscinum, Plagiomnium novae-zealandiae, Pohlia cruda, Polytrichadelphus magellanicus, Sphagnum australe, Thuidium furfurosum and Trichocolea mollissima.
Comments : The species name is in allusion to the more rigid plants, as compared to Trichocolea mollissima and T. hatcheri. Plants are mostly 1-pinnate, often relatively laxly so vs. the much denser 2-, 2–3-, or 3-pinnate branching of our other taxa. Trichocolea rigida is, also, a smaller species, with main axes mostly only 2–4 cm long; when dry it is a relatively dull plant, due to the conspicuously papillose surface, even terminal cells of cilia being clearly roughened (Fig. 25: 3, 5). The cilia are much less strongly tapered, with terminal cells ca. 4–5(5.5)× as long as in basal diam.; the septa between cells are never distinctly dilated (Fig. 25: 1, 3–5). In T. mollissima and T. hatcheri the cilia striking taper from base to apex, and are formed of much more elongated cells (to 15–19× as long as in basal diam.) and distal cilium cells are smooth or obscurely striolate.
