Cuspidatula monodon (Taylor ex Lehm.) Steph.
Anastrophyllum monodon (Hook.f. & Taylor) Steph., Hedwigia 32: 140. 1893.
Cuspidatula monodon (Hook.f. & Taylor) Steph., Sp. Hepat. 2: 126. 1901.
Jamesoniella monodon (Hook.f. & Taylor) N.Kitag., Bull. Nara Univ. Educ. 30: 47. 1981.
Type: Tasmania, 1824, Spence (“Hb. Greville et Taylor”).
Jungermannia geminiflora Colenso, Trans. & Proc. New Zealand Inst. 20: 243. 1888 (1887).
Type: New Zealand, North Is., Waipawa Co., S of Dannevirke, 1887, Colenso.
Jungermannia consimilis Colenso, Trans. & Proc. New Zealand Inst. 21: 47. 1889 (1888).
Type: New Zealand, North Is., Waipawa Co., S of Dannevirke, 1888, Colenso.
[Plate 12D; Fig. 158: 3, oil-bodies, p. 724; Figs. 159, 160]
Plants rigid, prostrate, the tips ascending, greenish brown to golden brown to purplish brown to reddish, nitid when dry, medium-sized, to 2.5 mm broad, rather short, seldom exceeding 2 cm long. Branching rather common, irregular, the branches all or in part ventral-intercalary, evidently from ventral ends of lateral merophytes (superficially appearing ventral in origin); Frullania -type branches rather common in some populations, apparently lacking in others, the half-leaf of similar form to the leaves, broadly connate with the first branch underleaf, which is ovate and broadly rounded at the summit. Stem wiry, with cortex in 1(2) layers of rigid, thick-walled cells, the exposed wall pigmented, notably thickened; medullary cells larger, strongly thick-walled with distinct, bulging corner thickenings. Rhizoids whitish, frequent, scattered. Leaves typically dorsally ± connivent in drying, when moist strongly dorsally assurgent grading to widely spreading, closely imbricate, 570–1100 µm wide × 750–1650 µm long, often adaxially concave, symmetrically to asymmetrically ovate-lanceolate, unlobed, apices caudate-piliferous, terminating in a uniseriate row of 8–15 cells, the distal cells highly elongated (to 8:1), otherwise entire, the tips often appearing hyaline and parched; dorsal margin arched and rather similar to the ventral in degree curvature (the leaves then symmetric) or the dorsal margin straight (the leaves then asymmetric), the ventral margin always arched, often sharply and locally incurved, entire but often a small blunt projection in distal portion of median third of margin, the base subcordate. Cells in median sector with strongly nodose trigones, the middle lamella conspicuous, 20–24 µm wide × 25–31 µm long; surface striate-papillose or smooth. Oil-bodies in all cells of leaf except distal cells of uniseriate row of leaf tips, occupying conspicuous portion of cell lumen, hyaline, glistening, 3(5)–(7)10 per cell, moderately botryoidal, subglobose and 4–6 µm in diam. to ellipsoidal, sometimes crescent-shaped and 5–7 × 9–12 µm. Chloroplasts large for cell size. Underleaves absent or vestigial, of 1–2 segments formed of 1–2 elongated cells, each ending in an ephemeral slime papilla; ventral merophytes appearing to be 2–3 cells broad. Asexual reproduction lacking (?or by fragmentation of leaf apices). Fungal partner absent.
Dioecious. Androecia on leading shoots, rather inconspicuous, becoming intercalary, often repeatedly so, ± spicate; bracts in 4–6 pairs, somewhat smaller and (when moist) more dorsally assurgent than leaves, the distal sector tapering and similar in form to leaves; lobule well developed, inflated, rather large for bract size, with a short, low wing, the free margin involute, distinctly long-acuminate distally; antheridia 1 per bract, the stalk biseriate, the stalk cells thick-walled; paraphyllia lacking. Gynoecia terminal on leading leafy shoots or relatively short gynoecial innovations; bracts large, sheathing the perianth, rather deeply concave, the insertion markedly narrow, the innermost pair of bracts often differing in lobe number, one bilobed, the other 3–4-lobed, in either case divided to 0.4–0.65, the lobes long-attenuate to acuminate to caudate-piliferous, terminating in a uniseriate row of up to 10 cells, the distal cells often elongated (to ca. 4:1), the lobe margins entire or with several often opposing sharp teeth and accessory lobule(s), the lamina on each side with a few small teeth and often an attenuated lobule; bracteole free or narrowly connate on 1 side, less than 0.55 width of bracts, distinctly narrow at the insertion, usually shorter than the bracts, oblong-lanceolate, acute or acuminate, undivided, on each side usually with a ciliiform to laciniiform to lobuliform process in distal half, the lateral processes never exceeding the length or size of the main, median process, the lamina also with a few small teeth and at times an accessory lobule below. Perianths frequent, large for shoot size, terete and ± inflated basally, the distal 0.6–0.7 deeply plicate, the plicae 5–7, rounded, separated by deep sulci, contracted at the often scorched mouth; mouth lobulate and ciliate/laciniate with stiff, piliferous cilia that terminate in a uniseriate row of 9–17 elongate, thick-walled cells, the lacinia bases often with a few, often opposed, stiff teeth.
Sporophyte as in generic description.
Distribution and Ecology : New Zealand: Auckland Islands (400 m), Antipodes Islands (200–300 m), Stewart Island (10–100 m), South Island (5–1525 m), North Island (60–1200 m), Chatham Islands (140–240 m); Australia: Tasmania, Victoria, New South Wales. Known from all ecological provinces of New Zealand. Extending to drier parts of some provinces, e.g., present in Marlborough (Pelorus Valley, Mt. Robertson), Banks Peninsula, Hawkes Bay and Wairarapa.
A common species throughout our area, but typically in rather exposed sites. It occurs on tree bark, e.g., on Nothofagus spp., Agathis australis (where particularly common), Phyllocladus, Weinmannia, Metrosideros, Coprosma and Libocedrus. Also on the bark of a number of species of Dracophyllum, e.g., in an area of rocky outcrops and shrub-heath communities including Coprosma foetidissima, Corokia buddleioides, Dracophyllum recurvum, Lepidothamnus laxifolius and Oreobolus pectinatus (exposed rocky ridge below summit area of “Little Moehau,” Mt. Moehau, Coromandel Forest Park, ca. 800–840 m). In “mesic conditions” also on old rotted, often decorticated logs. Commonly found on soil in rock crevices, particularly on exposed rock outcrops but also under forest. Also found on peat in penalpine bogs at sites subject to strong surface drying, and on soil, decomposing litter, and shrub bases in penalpine tussockland and scrub. Found with Campylopus introflexus, Hypnum cupressiforme, Jamesoniella colorata, Lepidozia obtusiloba, Leptotheca gaudichaudii, Pseudomarsupidium piliferum, Ptychomnion densifolium, Racomitrium crispulum, R. lanuginosum and R. pruinosum.
The species is strikingly tolerant of desiccation and may occur under quite rigorous conditions. For example, it was found on a thick soil layer over rock with Pseudomarsupidium piliferum over a lava flow within a forest of Metrosideros robusta (Rangitoto Island, 50–160 m). Also in the frost flats near Otupaka Stream (SW of Minginui, 700 m), a rigorous environment consisting of an open, low area with scattered Leptospermum scoparium and Dracophyllum subulatum with abundant Cladonia. And in the alpine zone it may form tight clumps on ice-eroded banks (below and W of Mt. Armstrong, SSW of Mt. Brewster, Mt. Aspiring Natl. Park, 1250–1450 m). The species may be a frequent component of the tree canopy bryoflora, and Allison and Child (1975, p. 94) mentioned that the species is “frequently an epiphyte, on branches rather than trunks.” Hodgson (1946, p. 73) indicated “K. W. Allison has a note to the effect that this species is sometimes found growing in the tops of tall forest trees.” Moreover, it may be found in fallen masses of bryophytes on the forest floor, e.g., in a mixed broadleaf forest dominated by Beilschmiedia and Vitex lucens, with Rhopalostylis sapida, Coprosma, Dacrycarpus dacrydioides and Hoheria (Kiwanis Reserve, N edge of Herekino Forest area, Northland, ca. 60–80 m). When found epiphytically, typical accompanying species are Acrochila biserialis, Dicnemon calycinum, D. semicryptum, Frullania aterrima, Holomitrium perichaetiale, Macromitrium longipes, Paraschistochila tuloides, Porella elegantula, Pseudomarsupidium piliferum, Pyrrosia eleagnifolia, Usnea rubicunda and Weymouthia cochlearifolia.
Comments : This species may be readily recognized in the field by the closely imbricate, unlobed leaves with the apex caudate-piliferous (Fig. 159: 1, 4, 5; Fig. 160: 1). It has been confused with Chandonanthus squarrosus, which can be a similar red or brown color, is found in similar habitats and has acute leaf lobe apices, but Chandonanthus has bifid leaves. The leaves, when moist, are often strongly dorsally assurgent (Fig. 159: 1), but may be widely spreading (Fig. 160: 1), and in some cases populations are either of one type of leaf orientation or the other. Leaves are typically dorsally ± connivent in drying. Plants are greenish brown to golden brown or reddish, prostrate, and often form pure, quite large patches. Underleaves are completely lacking, and ventral merophytes are very narrow (Fig. 159: 2). Schuster (2002a, p. 411, 413) was unable to document Frullania -type branches for the species. We have found that terminal branching is rather common in some populations of Cuspidatula monodon (Fig. 160: 1) while apparently lacking in others.
Spores are baculate and the bacula are truncate at the summit (Fig. 160: 8). This is yet another example of a predominantly corticolous hepatic having baculate spores (for a discussion see Engel and Gradstein, 2003).
