Calymperaceae

Shoots erect or creeping. Leaves linear, lingulate, ligulate or lanceolate; costa strong, incorporating stereids and guide cells and/or hyaline cells, often with a superficial layer of chlorophyllose cells; lamina in basal region of leaf well defined, largely unistratose, formed by large, empty, thin-walled, porose cells (cancellinae); in distal region of leaf composed of small, thick-walled, chlorophyllose cells (non-leucobryoid genera) or absent (most leucobryoid genera); leaf margin usually differentiated. Gemmae often produced on leaves, fusiform or filamentous, uniseriate. Dioicous. Sporophytes terminal, seta smooth; theca cylindrical; operculum rostrate or subulate. Peristome haplolepidous (16 teeth, usually papillose) and calyptra fugacious (Syrrhopodon, Leucophanes) or peristome absent and calyptra persistent (Calymperes). Spores papillose.

CALYMPERES Sw. ex F.Weber, Tab. Calyptr. Operc., 1813.

Shoots <0.5->3 cm high, sparsely branched, forming mats and tufts. Leaves dimorphic, gemmiferous and nongemmiferous. Nongemmiferous leaves 2-5 mm long, composed of a suberect, suboblong hyaline base narrowing slightly and abruptly into an erect to patent (incurved when dry), narrowly to broadly lingulate chlorophyllose limb that contracts abruptly to a broadly acute denticulate apex; costa ending immediately below apex, above hyaline base scabrid, internally incorporating stereids; chlorophyllose lamina plain to incurved, cells polygonal to rounded, mostly isodiametric to slightly longer than broad, <5-10(-12.5) x <5-7.5(-10) µm, ventrally subacutely protuberant, usually with a single papilla at the summit, dorsally flat and unipapillose; hyaline lamina sharply defined, usually occupying less than a third of the leaf length, cells dorsally and ventrally flat; leaf margin from leaf base to above apex of hyaline lamina with an intramarginal flattened band of long rectangular to linear thick-walled cells, distally, to near leaf apex becoming marginal and polystratose, in cross-section angled, superficial cells in surface view shortly subrectangular, many projecting acutely and towards leaf apex some forming small paired or triplet teeth; marginal lamina extending from leaf base to shortly beyond apex of hyaline lamina, in proximal leaf base formed by an entire, unistratose band of isodiametric to shortly subrectangular, thin walled, hyaline cells, mostly <3->7 cells wide; distally band becoming <7->12 cells wide with cells becoming smaller, thicker walled, chlorophyllose and forming a denticulate leaf margin. Gemmiferous leaves similar to non-gemmiferous leaves but abruptly narrowing into a linear leaf apex that consists largely of costa with a very narrow, recurved margin of chlorophyllose lamina; the latter becomes plain above to form a denticulate leaf tip. Gemmae produced in a bunch from the ventral surface of the costal apex, fusiform, uniseriate, pale reddish brown, smooth. Gametangia and sporophytes not seen in Ugandan material. [Sporophytes in West African specimens described by Edwards (1980).]

A pantropical species, broadly distributed within tropical continental Africa and the East African Islands.

Calymperes afzelii occurs at 900-1200 m a.s.l. on boles and buttresses of forest trees and on decaying logs.

Shoots reaching <0.5-2 cm high, often branched, forming dense mats and tufts; leaves 2-4 mm long, consisting of broad suboblong, suberect, semisheathing, hyaline base with an erect to spreading, narrowly to broadly lingulate (sometimes tapering) chlorophyllose limb that distally contracts to a broadly acute denticulate apex; costa with an inflated appearance, usually ending immediately below leaf apex but sometimes excurrent, largely scabrid, internally lacking stereids; chlorophyllose lamina plain to incurved, cells slightly longer than broad to slightly broader than long, >5-12.5(-14) x >5-10(-12.5) µm, ventrally acutely protuberant, dorsally unipapillose; hyaline lamina sharply defined, occupying over a third to half of leaf length, with cells at apex (hyaline and chlorophyllose) acutely exserted ventrally, in surface view appearing to overlap distally and laterally adjacent cells; leaf margin with intramarginal rib usually extending from leaf base to near apex (sometimes appearing marginal in proximal leaf base), in proximal hyaline leaf base formed by a flat, unistratose band of thick walled linear cells, distally becoming polystratose, composed of thick-walled shortly subrectangular to linear cells, some acutely protuberant at their distal ends; marginal lamina in mid hyaline base formed by a single row of narrowly subrectangular cells, mostly forming a tooth at their distal end; distally formed by a unistratose band of shortly subrectangular chlorophyllose cells, mostly 2-3 cells wide, outermost cells forming a dentate-serrate margin. Gemmae fusiform, smooth, pale reddish-brown, produced at leaf apex from dorsal and ventral surfaces of costa (sometimes from ventral surface of costa well below apex). Gametangia occurring frequently. Sporophytes frequent, exserted. Seta 5-7 mm long, reddish brown; theca 1-2 mm long. Spores (17.5-)20-25 µm diameter.

Frequently collected in Uganda, Calymperes pallidum is a pantropical species, widely distributed across continental Africa, Madagascar and the Seychelles.

This species occurs at 980-1550 m a.s.l. on decaying logs and stumps, on the trunks of tree ferns, palms and other trees.

Shoots erect, forming tufts and mats, often matted with rhizoids below. Leaves consisting of a largely hyaline, semisheathing base narrowing distally into a chlorophyllose limb; costa ending just below apex to shortly excurrent, smooth scabrid and/or papillose, internally with a single row of guide cells between dorsal and ventral bands of stereids, superficial cells often differentiated; laminal cells in chlorophyllose limb small and thick walled, in surface view polygonal to rounded, slightly longer than broad to slightly broader than long; flat or protuberant, smooth or papillose; laminal cells in hyaline base large, empty, with thin, porose walls, in surface view mostly subrectangular; leaf margin often bordered by a differentiated rib, entire, spinose, dentate, denticulate or ciliate. Gemmae often produced on leaves, uniseriate, fusiform or filamentous. Dioicous. Perichaetia and perigonia terminal (subperichaetial and subperigonial innovations often fertile), outer perichaetial and perigonial bracts similar to vegetative leaves, innermost perigonial bracts sheathing, oval and shortly pointed. Sporophytes exserted, seta smooth. Capsule cylindrical, sometimes with superficial stomata at base. Operculum with long, fine subulate beak. Peristome with 16 (usually ridged and/or papillose) teeth. Calyptra fugacious; narrowly mitriform, largely smooth and straw-coloured, with a reddened, slightly scabrous apex. Spores papillose.

Shoots reaching > 9 cm high, centipede-like, often sparsely branched, forming tufts and mats. Leaves <7-11 mm long, more or less in 3 ranks, consisting of a narrowly oblong, erect, semi-sheathing, hyaline base, abruptly narrowed into a linear, patent to spreading, chlorophyllose limb that, towards the apex, gradually tapers to a blunt, dentate tip; costa ending in leaf apex; ventral superficial cells shortly rectangular in surface view, mostly pluripapillose, distally some exserted as acute, spinose teeth; dorsal surface formed by stereids, distally with occasional groups of spinose teeth; chlorophyllose lamina slightly incurved; composed of subrectangular cells, isodiametric or longer than broad, (5-)7.5-12.5(-15) x 5-7.5(-10) µm, dorsally and ventrally pluripapillose, obscure; hyaline lamina occupying about a quarter of the leaf length, cells with strongly thickened dorsal walls (imparting slight orange tinge to lamina); marginal ribs strong, continuous from leaf base to apex; in leaf base a broad, largely unistratose band of thick-walled linear cells, mostly entire, sometimes distantly denticulate adjacent to distal hyaline lamina; distally becoming a broad, flattened polystratose band of stereids enclosing a horizontal layer of guide cells, some outermost cells of rib forming a single row of distant, acute teeth. Gemmae fusiform, mainly produced from ventral surface at apex of costa. Sporophyte exserted but often hidden among subperichaetial branches. Seta (rarely two from a perichaetium), about 8-11 mm long, dark red, apophysis with roughened surface, theca narrowly cylindrical, about 3 mm long, reddish-brown. Peristome teeth around 125–>160 µm long, brownish-orange, papillose. Spores c.12.5 µm diameter, finely papillose.

Syrrhopodon lamprocarpus is known only from Tropical Africa where it is widespread from São Tomé eastward to Uganda.

In Uganda it has been collected in forest between 1500 m and 1750 m a.s.l. on the trunks and exposed roots of trees, and on rotting stumps.

Syrrhopodon gaudichaudii Mont., Ann. Sci. Nat. Bot. sér. 2, 2: 376, 1834. (Fig. 4)

Syrrhopodon perdusenii W.D.Reese, Bryologist 97: 416, 1994. (Fig. 5)
(syn. Calymperes subdisciforme Dusén ex Besch.)

Syrrhopodon africanus (Mitt.) Paris, Index Bryol. 1244, 1898. (Fig. 6)

Syrrhopodon asper Mitt., J. Linn. Soc. Bot. 22: 303, 1866. (Fig. 7)
(syn. Syrrhopodon usambaricus Broth. ex S. Orbán)

Syrrhopodon gardneri (Hook.) Schwägr., Sp. Musc. Frond., Suppl. 2: 110, 1824. (Fig. 8)
(Syn. Syrrhopodon lisowskii S. Orbán)

Shoots reaching <1-5 cm high, often densely matted with rhizoids below, forming tufts and mats. Leaves 2-6 mm long, consisting of an erect, suboblong hyaline base narrowing into a broadly to narrowly linear-ligulate chlorophyllose limb that tapers near the apex to a narrow, blunt, dentate tip; costa ending immediately below apex in a dentate tip; superficial cells above hyaline base in surface view mostly shortly rectangular, usually some forming large acute teeth (sometimes arranged in distant, vaguely transverse rows); chlorophyllose lamina often incurved, sometimes strongly so; cells quadrilateral to rounded, slightly broader than long to slightly longer than broad, (5-)7.5-17.5(-20) x 5->10 µm, dorsally flat with one or two simple papillae, ventrally acutely protuberant with one or two simple papillae; hyaline lamina well defined, occupying from about a fifth to less than a third of the leaf length; marginal rib in hyaline leaf base formed by a unistratose band of thick-walled, linear cells, outermost row forming teeth, teeth adjacent to distal hyaline lamina prominent, acute, <15->20 µm long; above hyaline base becoming a polystratose rib, in cross-section with 3-4 angles (lacking stereids), superficial cells shortly subrectangular, some forming single, double or triple acute teeth at intervals along the margin (more frequent distally). Gemmae produced from the ventral (sometimes dorsal) surface of the costa, most often in a bunch at the leaf apex; fusiform, smooth, pale brownish green. Gametangia and sporophytes not seen in Ugandan collections.[Sporophytes of New World collections are described by Reese (1993).]

Frequently collected in Uganda, S. gardneri is a pantropical species, widespread across tropical continental Africa, and known from Madagascar and the Comores.

In Uganda, S. gardneri has been recorded between 1440-1685 m a.s.l. on buttresses and boles of forest trees, decaying logs, on rocks and soil in roadside banks and on the forest floor.

Syrrhopodon incompletus Schwägr., Sp. Musc. Frond., Suppl. 2: 119, 1824. (Fig. 9)

Shoots reaching 0.5->1.5 cm high, often sparsely branched, forming tufts and mats. Leaves >2-6 mm long, erect to patent when moist, incurved when dry; with a suboblong hyaline base slightly but abruptly narrowing into a linear to broadly linear-lingulate chlorophyllose limb, from above mid-limb tapering to a subacute, dentate apex (upper leaves tend to be relatively narrower than lower leaves); costa ending just short of leaf apex; dorsal and ventral superficial cells chlorophyllose, in surface view mostly shortly rectangular, some acutely protuberant, some forming long, multicellular, uniseriate, acute teeth (almost lamella-like, becoming more prominent in distal limb); chlorophyllose lamina plain with margins erect to incurved; cells in surface view polygonal to rounded, isodiametric or longer than broad, 5-12.5 x 5-7.5(-10) µm, dorsally and ventrally unipapillose or bipapillose; hyaline lamina well defined, occupying from about a fifth to less than a third of the leaf length; marginal ribs in proximal leaf base formed by unistratose band of thick-walled, linear cells; in distal hyaline leaf base absent (denticulate-dentate margin formed by chlorophyllose lamina, teeth projecting 7.5-10 µm from the leaf margin, intramarginal rib seen in New World collections not apparent in Ugandan material); above hyaline leaf base to near leaf apex formed by a polystratose rib of shortly subrectangular chlorophyllose cells, in cross-section subtriangular; some cells forming regular, usually paired or triplet, acute, lamella-like teeth (becoming more prominent in distal limb). Gemmae fusiform, papillose, produced in a small bunch at the leaf apex from the ventral and dorsal surfaces of the costa. Gametangia and sporophytes not seen in Ugandan material. [*Sporophytes of New World collections are described by Reese (1993).]

Newly recorded for Uganda (two collections from Kanyawara, Kibale National Park in Kabarole ), S. incompletus is uncommon in Africa, having been previously collected only in Tanzania and Zaire. This species is better known in the New World where it is represented by five varieties.

In Uganda, S. incompletus has been collected on tree boles and buttresses in forest at 1470 m and 1500 m.

LEUCOPHANES Brid., Bryol. Univ. 1: 763, 1827

Shoots erect, forming pallid tufts and mats, often matted with rhizoids below. Leaves consisting of a hyaline semisheathing base narrowing distally into a largely hyaline limb; costa thick, occupying whole of limb; composed of a unistratose median network of small, thick-walled chlorophyllose cells between a dorsal and a ventral layer of large, thin-walled, porose hyaline cells, and incorporating dorsally a central strand of stereids/substereids, strand often immersed, sometimes superficial and scabrid or smooth; lamina confined to basal region of leaf, unistratose, composed of large, thin walled, porose, shortly subrectangular hyaline cells; leaf margin bordered by a rib of stereids/substereids; entire, dentate or denticulate. Gemmae often produced on leaves, uniseriate, fusiform. Dioicous. Perichaetia and perigonia terminal (subperichaetial and subperigonial innovations often fertile), outer perichaetial and perigonial bracts similar to vegetative leaves but with proportionately shorter costa relative to hyaline lamina, innermost perigonial bracts sheathing, oval with an acute apex. Sporophytes exserted, seta smooth. Capsule cylindrical, with superficial stomata at base. Operculum subulate. Peristome with 16 (usually ridged and/or papillose) teeth. Calyptra fugacious; narrowly mitriform, largely smooth and straw-coloured, with a reddened, slightly scabrous apex. Spores papillose.

A single species, Leucophanes molleri, is recorded for Uganda.

Leucophanes molleri Müll. Hal., Flora 69: 285, 1886. (Fig. 10)

Shoots <0.5-2 cm high, simple or branching, densely leaved, forming pallid green tufts and mats. Leaves (2-)3-6 mm long, erect to reflexed, in cross-section ‘v’-shaped, basal region narrowly subelliptical tapering gradually into a narrowly lanceolate limb, apex acuminate-apiculate, rounded or truncate, often recurved, usually dentate or erose; costa broadening from base of leaf, distally tapering into limb and occupying almost entire width of leaf; cells of median chlorophyllose network viewed through the dorsal leaf surface mostly quadrilateral or pentilateral, slightly longer than broad to slightly broader than long, 6-12.5(-20) x 5-10 µm, in cross-section quadrate; superficial hyaline cells in surface view (sometimes in loosely transverse rows), mostly rectangular, mostly 37.5-80 x 7.5-12.5(->17.5) µm, (2-)5->8 times as long as broad, transverse walls mostly with a single round pore; strand of substereids, usually exposed at dorsal leaf surface, cells at dorsal angles of strand sometimes chlorophyllose and, towards the leaf apex, occasionally distally exserted as a small, acute tooth; marginal rib extending from the base of the leaf to apex, composed of thick-walled linear cells (substereids), in limb polystratose, and near leaf apex often giving rise to small, acute teeth. Gemmae produced in a bunch from the dorsal and ventral surfaces of the leaf apex; fusiform, uniseriate, smooth, pale green to brown. Gametangia and sporophytes not seen in Ugandan material. [Sporophytes described by Salazar Allen (1993).]

This species is widely distributed in Central America and tropical Africa, and has been frequently recorded in Uganda.

In Uganda, L. molleri has been collected between 1180-1710 m a.s.l., occurring on decaying logs and stumps, buttresses and trunks of trees, the trunks of tree ferns and lianas, and has also been discovered on rocks in roadside banks.

ACKNOWLEDGEMENTS

I am grateful to Miss J. Wilbraham for assistance in making up the plates, and to B.J. O’Shea for constructive comments on the manuscript for this paper.

References:

Eddy A. 1990. A handbook of Malesian mosses. 2. London: Natural History Museum.
Edwards S. 1980. A revision of West Tropical African Calymperaceae. 1. Introduction and Calymperes. Journal of Bryology 11 (1): 49-93.
Reese WD. 1993. Calymperaceae. Flora Neotropica 58: 1-102.
Salazar Allen N. 1993. Leucophanaceae. Flora Neotropica 59: 1-11.