Anastrophyllum (Spruce) Schiffn.
Anastrophyllum (Spruce) Steph., Hedwigia 32: 139. 1893.
Jungermannia subg. Anastrophyllum Spruce, J. Bot. 15: 235. 1876.
Lectotype (Schuster, 1961b): Anastrophyllum donianum (Hook.) Steph. (≡Jungermannia doniana Hook.)
Plants usually rather rigid, ascending to erect, especially if gemmae are formed, typically red to reddish brown to ± fuscous (even in shade), rarely deep green when in dense shade; minute and less than 500 µm wide to vigorous and to 3.5 mm or more wide; leafy shoots never becoming microphyllous and flagelliform distally. Branching sparingly, usually from lower sectors of shoots, the branches variable, at times lateral-intercalary, from middle or ventral ends of leaf axil; Frullania or Radula types sometimes present; dorsal-intercalary (Andrewsianthus -type) branching absent; geotropic, microphyllous, rhizoidous axes rare. Leaves firm, alternate, vertically oriented, the dorsal end of insertion transverse, extending to stem midline, often decurrent along dorsal midline of stem, the ventral half of insertion succubous (at very end of insertion again transverse or weakly decurrent), the leaves strongly concave, usually concave-canaliculate, the leaves bifid usually to 0.2–0.65; lobes acute to acuminate, rarely obtuse, the apices often somewhat incurved, the margins entire. Cells with large to coarsely nodose trigones that are confluent or separated by narrow thin-walled places (pits), the cell lumen often bounded mostly by the massive trigones, the cell walls sometimes uniformly thick-walled; cells in lobes and marginal sectors of lamina subisodiametric, those in a basal-median field usually conspicuously elongated, often 2–3:1; surface smooth or weakly (rarely strongly) papillose. Oil-bodies present in all cells, 3–4 per cell, granular to papillose, rather small. Underleaves lacking, sometimes even in gynoecial regions. Asexual reproduction usually lacking, at times present and by gemmae, the gemmae angular, formed at tips of upper leaves, sometimes on erect flagelliform shoots.
Dioecious (autoecious and paroecious in the Arctic Anastrophyllum sphenoloboides). Androecia ± spicate, the bracts ventricose at the base, the dorsal margin often with a distinct tooth toward the base; antheridia usually 1–2 per bract, the stalk 1–2-seriate; paraphyses absent. Gynoecial bracts usually bifid and similar to leaves except larger; bracteoles absent or present, free or connate on one side, unlobed, tapered to the summit. Perianths long-emergent, terete below, deeply plicate in distal sector, some taxa 3-plicate and with the dorsal face with a sulcus, some taxa pluriplicate, the perianths contracted toward the usually ciliate-dentate to ciliate-lanceolate, commonly bleached mouth.
Seta usually of the general type, some taxa ( subg. Crossocalyx) with 8(9) rows of outer cells and 4(5) rows of inner cells. Capsule wall usually 3–5-stratose, less often 2- or 2–3-stratose; outer layer of cells with nodular thickenings; inner layer of cells with semiannular bands, the bands often thin or incomplete on middle of tangential walls.
Spores papillose-short-vermiculate, spore:elater diam. ratio ca. 2:1.
Key to Species
One of the two largest genera in subfamilyLophozioideae (the other is Lophozia). We are accepting the broad delimitation given the genus Anastrophyllum by Schuster (1951, 1953, 1969c, 2002a), where the Sphenolobus, Crossocalyx and Eremonotus complexes are included within Anastrophyllum. Anastrophyllum contains ca. 30–35 species (Schuster, 2002a), but we note that Váňa (1984, p. 100) concludes, after further study, that there may be more (probably 35–40) species.
The genus Anastrophyllum has a broad range that includes both Arctic and south temperate species, but is concentrated in tropical-montane areas. Gradstein et al. (2001, p. 110), for example, stated that ca. 10 species occur at high elevations in tropical America and “ Anastrophyllum is one of the most characteristic genera of the páramo.” Ten species are known from the south temperate–subantarctic zone. Extra-New Zealand species are:
1) Anastrophyllum auritum (Lehm.) Steph. (type: South Africa, Table Mtn.) is pantropical. In Latin America a Cordilleran species that ranges from Mexico to Tierra del Fuego (Váňa, 1984); South Africa, Cape Prov. (Arnell, 1963; Váňa, 1982); Crozet Island (Grolle, 1971a); Marion Island (Grolle, 1971b). Váňa (1984, p. 102) states “this species, which is extremely variable ... has been described under many names (until now known as A. leucostomum (Taylor) Steph.).”
2) Anastrophyllum ciliatum Steph. occurs on South Georgia, Falkland Islands and the southern Magellanian region (see Engel, 1978, 1990a).
3) Anastrophyllum crebrifolium (Hook.f. & Taylor) Steph. (lectotype [fide Engel, 1978]: Chile, Prov. Magellanes, Cabo de Hornos, Hooker [FH!]); syn.: Anastrophyllum leucocephalum (Taylor) Steph. (type: Ecuador, Volcan Cayambe, 4265 m, Jameson [FH!, NY!]). The species occurs in southern South America (Tierra del Fuego, Patagonian Channel region), Juan Fernandez, higher elevations in Andes of Peru and Ecuador. See remarks in Engel and Braggins (1998).
4) Anastrophyllum involutifolium (Mont.) Steph. (syn.: Anastrophyllum decurvifolium (Sull.) Steph.) of southern South America (Tierra del Fuego, southern Patagonian Channel region) and Juan Fernandez (see Engel, 1978).
5) Anastrophyllum minutum (Schreb. ex Cranz) R.M.Schust. (type: Greenland) is a broadly distributed Holarctic species (cf. Schuster, 1969c) and occurs in the Southern Hemisphere as a disjunct at high elevations in New Guinea and South Africa (Arnell, 1963 as Sphenolobus minutus (Schreb. ex Cranz) Steph.; Váňa, 1982). Reported from Kerguelen by Hooker and Taylor (1844a) and Gottsche et al. (1844–47); the record is probably erroneous (Váňa, 1982).
6) Anastrophyllum piligerum (Nees) Spruce (type: Java), syn.: Anastrophyllum incumbens (Lehm. & Lindenb.) Steph. (type: Tristan da Cunha). A pantropical species, including presence in Zimbabwe on Inyanga Mtn., and Tristan da Cunha (Váňa, 1982, 1984).
7) Anastrophyllum tristanianum J.J.Engel is endemic to Tristan da Cunha (see also comments in Engel and Braggins, 1998).
The generic description after Schuster (2002a) but modified.
References: Engel and Braggins (1998); Schuster (1966c, 2002a).