Pseudomarsupidium piliferum (Steph.) Herzog ex Grolle
Marsupidium piliferum Steph., Sp. Hepat. 3: 386. 1908.
Lectotype (fide Grolle, 1972a): Australia, New South Wales (“Australia felix”), Ferd. von Müller (G!); syntype: Chile, Corral, Krause, ex herb. Gottsche (G!).
Tylimanthus bisetulus Steph., Sp. Hepat. 6: 246. 1922.
Adelanthus bisetulus (Steph.) Grolle, J. Hattori Bot. Lab. 35: 359. 1972.
Lectotype (fide Grolle, 1972a): New Caledonia, in jugo Dogny, 1040 m, Lerat 361 (G!).
Adelanthus humilis Steph., Sp. Hepat. 6: 446. 1924.
Type: New Caledonia, in jugo Dogny, 1040 m, 1909, Lerat 207 (G!).
[Plate 12C; Fig. 128: 5, oil-bodies, p. 558; Figs. 129, 130]
Plants chestnut-brown, dull, the shoot system consisting of prostrate, copiously branched, basal stolons that are often partially embedded in the substrate, and erect, leafy shoots. Branching always ventral-intercalary, for the most part from basal sectors of plant, the stolons with small leaves with apices broadly rounded and devoid of spines and underleaves that are minute, inconspicuous and irregularly acute. Stems of erect shoots narrow for plant size, stiff and wiry, chestnut-brown; stem in cross section with cells ± uniform in size, the cortex in 1(2) layers, reddish brown, with thickened walls. Rhizoids copious, on all sides of stolons, lacking entirely from leafy sectors of erect branches. Leaves of erect shoots stiff, brittle, fragile, reduced and scale-like below, becoming densely imbricate, concave, not vittate, the orientation ± transverse to succubous, the insertion narrow, oblique and slightly recurved at ventral end, the leaves ovate to orbicular to feebly oblate, abruptly narrowed at the base; apex with 2(rarely 3), elongated, often ventrally bent or flexed, hyaline or pale brown awns (on suboptimal shoots reduced to thorns), the dorsal awn at juncture of apex and dorsal margin and typically appearing as a continuation of the margin, the awns consisting of a uniseriate row of to 9(10) cells, the awn cells elongated, 17–30 µm wide × (65)72–114 µm long (typically 2.4–6.6:1), with strongly thickened walls, the terminal cell sharply pointed, the apex and margins otherwise entire; dorsal margin often erect, inflexed and ± clasping the stem, particularly when dry, the base broadly decurrent by a distinct acute strip, ± straight to broadly and gently curved; ventral margin broadly ampliate. Cells with thin walls, with medium to weakly knot-like trigones; subapical cells isodiametric, 24–34 µm wide and long; median cells nearly isodiametric, 26–38 µm wide × 30–38 µm long; median-basal cells larger, not markedly elongated; marginal row of cells with exposed wall strongly thickened, the “half-trigones” along margin very large. Oil-bodies occupying a conspicuous portion of the cell lumen, often appearing packed, distinctly greyish, 10–19 per median leaf cell, coarsely papillose, the spherules soon swelling and coalescing and the oil-bodies then increasing botryoidal, many oil-bodies globose, some elliptic, notably variable in size within a cell, some ± 3–4× smaller than others, some of larger ones at times with a crease or depression, the oil-bodies 5.8–7.7 × 10.1–12 µm, to 8–8.5 × 11–16 µm to 10–11 × 11–13 µm, spherical ones 4.8–8.2 µm in diam.
Androecia on abbreviated branches from stolons or bases of erect, leafy branches, pale, compactly spicate, often clustering, often coiled, often somewhat dorsiventrally compressed; bracts in 4–6 pairs, strongly and deeply saccate, the dorsal margin entire, the apex entire or irregularly crenulate, the marginal cells elongated at right angles to margin; antheridia 1 per bract, the stalk rather long, uniseriate; bracteoles present at least in basal half of androecium, scale-like, irregular in shape, the apex and often margins rather coarsely crenulate. Gynoecia similar in position to androecia, bulbous, fleshy and rhizoidous at base; bracts in 2–3(4) series, pale, the innermost (non-perianth-forming cycle) free from one another, conspicuously larger than those below, the apex irregularly spinose dentate-lobulate, the margins of the lamina dentate by 1–2-celled teeth, the terminal cell rather elongate, the lamina often subentire below; bracteoles subequal to bracts or somewhat smaller, at times much more so, at times reflexed; innermost series of bracts and bracteole (the perianth-forming cycle) extremely variable, at times united and developed as a distinct perianth, at times ± deeply divided and only basally connate, at times completely divided into separate, free bracts; perianth (when present) short, obscurely to rather distinctly trigonous, variable in shape, narrowly to broadly and stoutly ovate-cylindrical, with sharply lanceolate scales inserted on basal sector, often plicate, straight or more often narrowed toward the mouth, the mouth broad to narrow and contracted, the mouth (or, in absence of a perianth, the apex of the perianth-forming cycle of bracts) irregularly and densely spinose-ciliate, the cilia composed of a uniseriate row of 3–6 elongate, often thick-walled cells. Mature calyptra thin, membranous, 3–5-layered medially, collapsing distally after capsule dehiscence, extending to level of perianth mouth (when perianth present) or beyond, with sterile archegonia at calyptra base or if elevated then remaining in basal third, a few paraphyllia sometimes present at calyptra base (distal to bracteole).
Seta very long (to 3.5 cm long), seen only in collapsed state. Capsule ellipsoidal, the wall 54–64 µm thick, of 4–5 layers; outer layer of cells quadrate to short-rectangular, with imperfect two-phase development, the longitudinal walls with strong nodule-like thickenings alternating with primary walls either devoid of thickenings or with a few nodular thickenings, the transverse walls with thickenings absent or, often, present; innermost layer of cells for the most part irregularly narrowly rectangular, with semiannular bands, the bands narrow, sometimes incomplete, often forked and sometimes anastomosing to delimit local fenestrae, the longitudinal and transverse walls at times with nodule-like thickenings.
Spores 32.2–37.4 µm in diam. (including bacula), yellow-brown, baculate, the bacula rather widely spaced, slightly wider at base, truncate at the apex and often flared, the ground surface otherwise with a network of delicate, fine, narrow, rather distantly spaced low ridges, the ridges sporadically forked, the surface of ridges without papillae; spore:elater diam. ratio 3.5–4.1:1. Elaters somewhat adhering to inner face of capsule wall after capsule dehiscence, tortuous, 9.1–10.6 µm in diam., bispiral to tips or the ends at times somewhat attenuated and in that sector unispiral, the bands 2.9–3.4 µm wide.
Distribution and Ecology : New Zealand: Campbell Island (60 m), Auckland Islands (275–400 m), Stewart Island, South Island (180–1250 m), North Island ([50]300–1100 m), Chatham Islands; Australia: Tasmania, Victoria, New South Wales; New Caledonia, southern Chile, Juan Fernandez, Inaccessible Island, Nightingale Island. In New Zealand known from Fiordland, Westland, Western Nelson, Volcanic Plateau, Taranaki, Auckland (Mt. Moehau, Great Barrier Island) and Northland EPs.
In New Zealand occasional but not frequent. In the South Island in forests of Nothofagus solandri var. cliffortioides or of N. menziesii forest where it may form thick, often pure, mats over thin soil on cliff faces or on rock ledges of outcrops, or, less often, on bark of Nothofagus. In the North Island in scrubland or areas with low open stunted trees and shrubs, often in fairly exposed sites. It occurs epiphytic on trunks of Kunzea ericoides (with Hymenophyllum and Bazzania hochstetteri), or of Weinmannia racemosa, or (Mt. Moehau, ca. 840 m) on upper branches of stunted Phyllocladus glaucus in an area with stunted W. silvicola, Coprosma foetidissima and Corokia buddleioides or (Mt. Te Aroha, 950 m) on small upper branches of Weinmannia in a stunted mixed-age N. menziesii, Weinmannia and Ixerba brexioides forest. Other known hosts are Brachyglottis rotundifolia, Leptospermum scoparium, Mida cunninghamii, Neomyrtus pedunculatus and Prumnopitys taxifolia. In exposed sites tufts of the species are often a dull olive-brown or chestnut-brown. Also on ground at cliff bases or on weathered rock, at times associated with Jamesoniella colorata and Cuspidatula monodon. Also at a considerably lower elevation (50–160 m) on Rangitoto Island; the species occurred on a thick layer of soil over shaded rock in a Metrosideros forest developed over a lava flow. At Stockton Plateau it occurs in soil pockets in deeply eroded sandstone pavements among low Leptospermum scoparium and Rytidosperma setifolium, with Acromastigum anisostomum, Campylopus clavatus, Cladonia confusa, Dicranoloma robustum, Holomitrium perichaetiale, Jamesoniella colorata, Rhacocarpus purpurascens, Siphula decumbens and S. fragilis. At other sites on soil or rock it has also been found with Anastrophyllum schismoides, Campylopus introflexus, Herbertus oldfieldianus, Hypnum chrysogaster, Lepicolea attenuata and Racomitrium pruinosum. Where epiphytic it has been found with Bunodophoron scrobiculatum, Chandonanthus squarrosus, Cuspidatula monodon, Dicnemon calycinum, Herbertus oldfieldianus, Heteroscyphus menziesii, Holomitrium perichaetiale, Jamesoniella colorata, Lepicolea attenuata, Leptostomum macrocarpum, Paraschistochila tuloides and Plagiochila circinalis.
Comments : This strongly fuscous plant has ± transversely to succubously oriented, concave leaves with the dorsal margin stiffly elevated, and the leaf apex with 2 hyaline, stiff awns (rarely reduced and tooth-like). It is not likely to be mistaken for anything else, although it somewhat resembles species of Marsupidium in aspect, but these are green, and never have the blackish brown color of the Pseudomarsupidium.
The gynoecium of Pseudomarsupidium piliferum is variable in the extent of perianth development. The uppermost gyre of bracts and bracteole (the perianth-forming cycle) may be a) fused nearly to the summit, thus forming a distinct and well-defined perianth (Fig. 129: 7); b) connate only at the base, the perianth lobes thus being deeply divided (Fig. 129: 6); or c) completely free, a true perianth thus completely lacking (Grolle, 1972a, pl. 10 n). The calyptra is 3–5-stratose in the median sector and may be within the confines of the perianth (Fig. 129: 7) or exceed it (Fig. 129: 8; Fig. 130: 1). Figures 129: 7 and 8 were redrawn from Grolle (1972a), who based them on the same collection (Schwabe 53a from Chile). Engel 16275 from Tasmania has a well-developed perianth and a calyptra that extends to the perianth mouth. Shoots of Braggins 99269A (Mt. Taranaki, AK) have sporophyte-bearing gynoecia with the cycle of all 3 perianth-forming bracts fused to one another only at the extreme base, but are otherwise free above (Fig. 130: 1). The perianth-forming bracts are deeply concave to inflated-pouched, are weakly to distinctly longitudinally plicate (Fig. 130: 1) and are similar in appearance to plicate perianths in various other hepatic families. The perianth-forming bract apex has cilia with a uniseriate row of 3–6 elongate cells (Fig. 130: 3). Unfertilized archegonia are positioned at or near the base of the calyptra (Fig. 130: 1).
Pseudomarsupidium piliferum is yet another species exhibiting a suite of sporophyte and spore characteristics shared by unrelated taxa that are uniformly or mostly corticolous. For a discussion see Engel and Gradstein (2003).
