MALOU - Lophoziaceae

Mosses and Liverworts of Uganda

Lophoziaceae

By J. Vána (illustrations by M. C. Watling)

Plants small to large, green, yellowish to brown or blackish, sometimes with red colour. Shoots creeping, prostrate, suberect, ascending to erect, simple or mostly with lateral-intercalary, rarely terminal branching, in some species microphyllous flagelliform branches present. Stems soft-textured to rigid, mostly with differentiated cortex with thick-walled cells. Rhizoids scattered, rare or copious. Leaves succubous, mostly obliquely, rarely transversely inserted, mostly distant to imbricate, concave to canaliculately concave, predominantly 2-lobed (and obtuse to acute), but also unlobed or 3-4-lobed leaves occur; margins mostly edentate, sometimes also dentate to ciliate; lobes predominantly triangular to ovate-triangular, equal, subequal to unequal, sinus acute to lunulate. Underleaves absent or present, lanceolate to 2-lobed. Cells mostly collenchymatous with large to nodulose trigones, rarely thick-walled and/or without trigones. Cuticle smooth to papillose. Oil-bodies 2-6 to numerous, granular, mostly not segmented. Gemmae sometimes present, ovate to angulate, 1-2-celled, mostly on modified or unmodified apices of shoots or apices of leaf lobes, green, yellowish-green, sometimes reddish to vinaceous. Monoicous (autoicous, paroicous to heteroicous) or more commonly dioicous species. Androecia terminal, becoming intercalary or subgynoecial, male bracts mostly similar to leaves, saccate. Female bracts also similar to leaves, sometimes larger to more lobed, free or connate; innovations under gynoecia sometimes present. Bracteolae present or lacking. Perianth emergent, mostly cylindrical, pyriform to ovoid, apex contracted to entire, dentate to ciliate, rarely tubular mouth; always without perigynium. Seta mostly from unspecialized cells. Capsule ovoid, ellipsoidal to globose, with 2-5 (-8)-stratose wall; elaters mostly 2-spiral.

1.	Leaves deeply 3- or 4-lobed to 0.8-0.9 their length. Underleaves large, to 0.8-0.9 bifid	2
	Leaves not deeply (mostly up to 0.5 their length or less) 2- or 3-lobed. Underleaves absent or only small, lanceolate to subulate	3
2.	Leaves asymmetrically 3(-4)-lobed, ± obliquely inserted, succubous, with sharp marginal teeth; cells collenchymatous, with large to nodose trigones	*Plicanthus*
	Leaves ± symmetrically (3-)4-lobed, ± transversely inserted, with only basal teeth or laciniae; cells thick-walled, with poorly defined or small trigones	*Tetralophozia*
3.	Leaves 3-lobed (exceptionally 2-lobed); leaf lobes clearly unequal	*Tritomaria*
	Leaves 2-lobed (exceptionally 3-lobed with small third lobe); leaf lobes equal or rarely subequal	4
4.	Plants with Anomoclada-type branches producing stoloniform, flagelliform or microphyllous branches	*Andrewsianthus*
	Plants without Anomoclada-type branches (if these branches very rarely present, never producing flagelliform or microphyllous branches)	5
5.	Leaf insertion mostly transverse (excl. decurrent part); plants mostly brownish to fuscous	*Anastrophyllum*
	Leaf insertion oblique, succubous; plants mostly green to yellowish green	*Lophozia*

Andrewsianthus R.M.Schust., Rev. bryol. lichénol. 30: 66, 1961

Plants small to medium sized, green to brownish. Shoots prostrate to suberect, typically with lateral-intercalary branching Anomoclada-type, producing flagelliform, microphyllous attenuate geotopic or plagiotropic stolons; terminal branching only exceptional. Stems rigid, with 1-2 rows of cortical thick-walled cells. Leaves succubous, obliquely to nearly subtransversely inserted, mostly 2-lobed, but also species with unlobed, retuse or 3-lobed leaves exist; leaf margin mostly entire. Underleaves absent or only small, reduced. Cells mostly with distinct, often nodulose trigones. Cuticle mostly papillose or striolate. Gemmae totally absent. Dioicous. Androecia spicate, male bracts saccate, mostly with 1-2 basal teeth, 1-androus. Gynoecia terminal, female bracts larger than normal leaves, sometimes toothed; bracteoles mostly distinct. Perianth emergent, multiplicate, contracted to a dentate, lobulate or laciniate mouth. Capsule oval, mostly with 3-layered wall. Spores mostly verrucose to minutely spinose.

Andrewsianthus bilobus (Mitt.) Grolle, Trans. Brit. Bryol. Soc. 4: 437, 1963

Plants medium sized, about 2 cm long and 1.1-1.8 mm wide, mostly brownish, rarely deep green. Shoots prostrate to ascending, simple or with lateral-intercalary branching, producing flagelliform, partly microphyllous stolons. Stem with 1-2 rows of cortical thick-walled cells. Leaves succubous, obliquely inserted, slightly concave, distant to imbricate, circular to slightly reniform, up to 0.3 of the length 2-lobed, sinus acute to subacute. Lobes widely triangular, acute to subapiculate, with entire margins. Underleaves only at apices of shoots, highly reduced up to papillae, mostly totally absent. Cells thin-walled, with large to nodose trigones, marginal 15-20 µm, median 20-25 µm, basal up to 30-35 µm. Cuticle striolate. Dioicous, very rarely fertile. Perianth ovate, plicate at apex.

Known from the Rwenzoris and Mt. Elgon as an epiphyte or growing on wet rocks in subalpine forests. Also known from Bioko, Cameroon, D.R. Congo, Ethiopia, Rwanda and Tanzania.

Anastrophyllum (Spruce) Steph., Hedwigia 32: 139, 1893

Plants small to large, from 0.5 to about 10 cm, mostly brownish to fuscous, reddish or blackish, rarely green. Shoots rigid, prostrate, ascending to erect. Stems with firm cortex and collenchymatous medullary cells; simple or lateral-intercalary, terminally to rarely ventral-intercalary branched. Leaves sometimes distichous, vertically oriented, transversely or subtransversely inserted, but with ventral base succubous and sometimes decurrent, constantly 2-lobed to 0.1-0.6 their length. Underleaves absent. Cells thin- or rarely thick-walled, with small to large, nodulose to confluent trigones. Cuticle smooth to striolate. Gemmae mostly absent. Mostly dioicous, rarely autoicous or paroicous. Androecia spicate, male bracts saccate, with basal teeth, 1-2-androus. Gynoecia terminal, female bracts similar to leaves, 2-lobed to 4-lobed. Perianth emergent, in terminal part sometimes decolourate, large, at least distally pluriplicate, contracted to dentate to ciliate mouth. Capsule wall with 2-5 layers. Spores mostly papillose, 10-15 µm.

	Leaves widest near the base, the ampliate dorsal base extended across the stem; cells with trigones, basal cells elongated; gemmae lacking	*A. auritum*
	Leaves not extended across the stem; cells thick-walled, lacking trigones; gemmae present	*A. minutum*

Anastrophyllum auritum (Lehm.) Steph., Spec. Hep. 2: 120, 1901

Plants in deep brown to reddish loose to compact patches. Shoots about 0.5-4 cm long and up to 0.5-1.0 mm wide. Stems prostrate with ascending tips, suberect to erect, simple or mostly branched; cortical cells in 2-3 layers deep brown, thick-walled. Leaves contiguous to imbricate, obliquely succubously inserted with antically decurrent ampliate base and mostly with basal appendages on antical margin, erect-spreading, concave, ovate to circular, 2-lobed to 0.25-0.5 their length. Leaf lobes acute to subacute, strongly incurved, sinus acute to subacute, somewhat gibbous. Basal appendages acute, subulate, up to 4-6 cells wide and 10-15 cells long, sometimes smaller to nearly absent in poorly developed plants. Underleaves lacking. Cells with irregularly thickened walls and large to bulging trigones, 10-15 µm on the leaf apex, 15-20 µm in mid-leaf, basal cells elongated up to 30 µm. Gemmae lacking. Dioicous. Male bracts strongly saccate, 1-2-androus. Female bracts similar to leaves, larger, sometimes with some blunt teeth on the margin and gibbous sinus. Perianth subcylindrical, in upper part often decolorate, pluriplicate near the ciliate mouth.

Widely distributed and common in subalpine forests (from 2870-4450 m) of the Rwenzoris, Mt Muhavura and Mt Elgon, growing on rocks and as an epiphyte on trees. In Africa also known from Bioko, Cameroon, D.R. Congo, Ethiopia, Kenya, Lesotho, Malawi, Rwanda, South Africa and Tanzania.Widely distributed and common in subalpine forests (from 2870-4450 m) of the Rwenzoris, Mt Muhavura and Mt Elgon, growing on rocks and as an epiphyte on trees. In Africa also known from Bioko, Cameroon, D.R. Congo, Ethiopia, Kenya, Lesotho, Malawi, Rwanda, South Africa and Tanzania. Widely distributed in mountains of Neotropics from Mexico to Fuegia, also known from Borneo, New Guinea and some subantarctic islands (Crozet Is., Marion I.).

Anastrophyllum minutum (Schreb.) R.M.Schust., Amer. Midl. Naturalist 42: 576, 1949 (see figure above)

Plants in deep green to fuscous patches or admixed with other bryophytes. Shoots small to medium sized, 1-3 cm long up to 1.2 mm wide. Stems prostrate or ascending, simple or with sparsely lateral-intercalary or terminal branching; cortical cells hardly differentiated, thick-walled. Leaves distant to imbricate, pectinately oriented, widely ovate to subquadrate, canaliculately concave, at least in the dorsal part transversely inserted, 2-lobed to 0.2-0.4 their length. Leaf lobes triangular, acute to apiculate, sinus subacute to acute. Underleaves absent. Cells subquadrate, with thick walls and nearly absent trigones, 12-20 µm in leaf lobes, 15-25 µm in mid-leaf, not elongated at the base of leaf. Cuticle faintly verrucose. Oil-bodies 2-6 per cell, granulose. Gemmae on the apices of lobes of reduced leaves and on the stem apex, mostly angular, 2(-4)-celled, reddish. Dioicous, mostly sterile. Male bracts concave, 1-2-androus. Female bracts larger than leaves, sometimes 3-4-lobed, the lobes entire. Perianth emergent, subcylindrical, with plicate upper part, mouth dentate. Seta of many cell rows, capsule wall 2-3-stratose. Spores 10-15 µm, brown.

Collected only twice: on NW slope of Mt Muhavura, on a moist, steep rock in a ravine, at 4000 m and on Mt. Elgon, in tufts in the crevices of the volcanic rocks. Also known from Cameroon, D.R. Congo, Kenya, South Africa and Tanzania. Widely distributed in the holarctic region, southernmost to Colombia, Himalayas, Borneo and New Guinea.

Lophozia (Dumort.) Dumort., Recueil d´Observ. Jungerm., p. 17, 1835

Plants very variable, small to large, green to brownish, sometimes reddish to vinaceous. Shoots mostly prostrate to suberect, simple or with very variable branching; mostly lateral-intercalary, but often terminal to ventral-intercalary, flagelliform branches nearly absent, but leafless stolons sometimes present. Leaves 2-4-lobed, obliquely inserted and succubous, mostly concave. Underleaves sometimes present. Cells also very variable, from thin-walled to strongly collenchymatous with large to nodulose trigones. Gemmae mostly present, ovate to angular, mostly 1-2-celled. Dioicous, autoicous or paroicous plants. Androecia spicate, terminal or intercalary, with saccate male bracts. Female bracts mostly similar to leaves, larger, at least ± more-lobed. Perianth cylindrical to clavate, emergent, at least in upper part plicate. Capsule wall 3-5-layered.

1.	Leaves with spinose to dentate margins; plants with leafless stolons and ventral-intercalary and also Anomoclada-type lateral branches; gemmae yellowish-green	*L. jamesonii*
	Leaves with entire margins; plants without leafless stolons, ventral-intercalary and also Anomoclada-type branching	2
2.	Cells equally thick-walled; plants very compact, julaceous, with closely imbricate leaves, gemmae reddish	*L. decolorans*
	Cells thin-walled. Plants with remote, not closely imbricate leaves, gemmae violet	*L. excisa*

Lophozia decolorans (Limpr.) Steph., Spec. Hep. 2: 128, 1901

Plants very compact, yellowish to yellowish brown, often with decolourate distal parts. Shoots 1-2 cm long, rigid, creeping, simple to sporadically branched. Leaves closely imbricate, concave, very broadly orbicular to ovoid-orbicular or reniform, 0.8-1.0 mm wide and 0.5-0.7 mm long, narrowed to shallow lobes that are not widely separate, sinus descending to 0.1-0.15 (-0.2), lunate. Lobes ± subequal, much broader than long, apiculate. Underleaves lacking. Cells ± thick?walled, 15-25 µm in the margin, 20-30 ×15-25 in the middle of leaf. Gemmae reddish, 1-2-celled, 4-5-angled, quadrate to subquadrate, 20-25 µm. Dioicous. Androecia terminal, becoming intercalary, with 3-6 pairs of less imbricate male bracts, bracts monandrous. Gynoecia terminal, female bracts reniform, 2-lobed, lobes apiculate. Perianth decolorate above, shallowly plicate, mouth crenulated with 1-celled teeth. Spores 12-15 µm.

Collected only twice in Mt. Elgon area on isolated shrubs of Erica trimera on basalt plateau at 3460-4000 m. Very disjunct holarctic species (Western Canada, Alps, Bulgaria, Norway, Russian Arctic, Himalaya), in Africa also known from Cameroon, D.R. Congo and Tanzania.

Lophozia excisa (Dicks.) Dumort., Recueil d´Observ. Jungerm., p. 17, 1835

*Plants* pale green, intermixed among other bryophytes. Shoots 2-3 cm long, flaccid, prostrate. Stems with thin-walled cortex, simple or lateral-intercalary branched. Rhizoids relatively common. Leaves remote, flaccid, obliquely inserted, succubous, nearly circular, subequally 2-lobed, rarely 3-lobed with dorsal lobe smaller; sinus obtuse, slightly gibbous; lobes triangular, subacute. Underleaves absent. Leaf cells thin-walled with small trigones, marginal 20-25 µm, median 25-30 µm, basal up to 35-40 µm. Cuticle smooth. Gemmae violet, polygonal. The known African specimens are sterile. Paroicous, with hypogynous somewhat saccate male bracts in 2-6 pairs. Perianth ovate, emergent. Collected only once in the Rwenzoris, Mijusi Valley, among mosses under “Alchemilletum” on dry moraine ridge at 3900 m. Widely distributed holarctic species with bipolar range, common also in antarctic and subantarctic region; in Africa known probably also from Lesotho. On the basis of the tendency to produce 3-lobed leaves, its cross section of stem, cell size and the form of typical gemmae it seems that L. hedbergii is only a modification of L. excisa, until now regarded as a bipolar species. There are no unambiguous differences from L. hedbergii; African plants of L. excisa represent mod. laxifolia.		a, shoot; b-f, leaves; g, mid-leaf cells; Hodgetts 3109 (Herb. N.G. Hodgetts) , U.K., Norfolk.

Lophozia jamesonii (Mont.) R.M.Schust., Austral Hepaticae 2: 279, 2002

Plants pale green, yellowish to brownish, in pure patches or intermixed among other mosses. Shoots 0.4-1.5 cm long, prostrate to ascending, simple or rarely lateral-intercalary (Plagiochila- and Anomoclada-type), exceptionally ventral-intercalary branched; the last one branching producing leafless stolons. Stems with relatively thin-walled cortical cells. Rhizoids relatively frequent, colourless. Leaves slightly obliquely inserted, succubous, distant to imbricate, forming a crispate head, erect-spreading, oblong to obtrapezoidal, subequally 2-lobed or rarely 3-lobed, postical lobe the largest; sinus descending to 0.3-0.5 their length; at least upper leaves with spinose margins, basal leaves sometimes with subentire margins; lobes ending with spinous tooth. Underleaves sometimes present, subulate to reduced to 1-2 cells or totally absent. Cells thin-walled, with small or large, but not bulging trigones, marginal cells 22-30 µm, median ones 25-30 × 30-45 µm, basal cells elongated to 55 µm. Cuticle smooth or nearly so. Gemmae stellate, mostly 1-celled, 11-15 µm, produced on apices of the upper leaves. Autoicous or dioicous (? pseudodioicous). Androecia spicate, terminal, becoming intercalary, bracts in 5-12 pairs, with toothed margins, monandrous. Gynoecia terminal, female bracts larger than leaves and somewhat crispate, less deeply lobed and more spinose-dentate; bracteole		a, shoot; b-j, leaves; k, mid-leaf cells; Herb. J. Vana. TANZANIA, Kilimanjaro.

present, lanceolate to triangular, not connate with bracts. Perianth cylindrical to pyriform, plicate in upper part, mouth ciliate-dentate with teeth 1-3 cells long. Capsule with 2-stratose wall. Spores 15-18 µm.

Collected only twice in the Rwenzoris, Mijusi Valley, on moist crevice of a steep rock, 3900 m and in NW slope of Mt Muhavura, on moist rock in a ravine, 4000 m. Also known from D.R. Congo, Kenya, Rwanda, Tanzania and from Costa Rica, Panama, Venezuela, Colombia, Ecuador and Peru.

Plicanthus R.M.Schust., Nova Hedwigia 74: 484, 2002

Plants in loose patches, mostly yellowish to golden brown. Shoots stiff and rigid, relatively large, 3-12 cm long and 1.0-5.0 mm wide. Stems procumbent or erect, simple or sparsely lateral-intercalary branched, with 2-4 rows of cortical cells, medullary cells larger, also stellate and firm-walled; ciliate paraphyllia mostly present. Leaves obliquely inserted, succubous, asymmetrically 3(-4)-lobed up to 0.8-0.95 of their length, abaxially concave to sulcate; leaf lobes narrowly ovate to lanceolate, concave to sulcate, with dentate to ciliate margins and acuminate apices. Underleaves small, less than 0.5 of the size of leaves, to 0.8-0.9 2-lobed, the lobes similar to leaf lobes. Cells thin-walled, with large, coarse, sometimes confluent trigones, basal cells somewhat elongated. Cuticle smooth to striate. Gemmae absent. Dioicous. Androecia unknown. Gynoecia terminal, female bracts similar to leaves, slightly larger, with ciliate to laciniate margins, connate with bracteoles; bracteoles 2-lobed. Perianth longly emergent, 5-6-plicate to base, with ciliate to spinose teeth at mouth. Seta massive. Capsule with 4-stratose wall. Spores papillose.

	Large plants up to 10-12 cm long; stems with paraphyllia; leaf lobes with 15-20 cilia on each margin	P. giganteus
	Smaller plants up to 3-4 cm long; stems without paraphyllia; leaf lobes mostly with 5-7 cilia on each margin	P. hirtellus

Plicanthus giganteus (Steph.) R.M.Schust., Nova Hedwigia 74: 486, 2002

*Plants* in loose yellowish to brownish patches. Shoots rigid, 10-12 cm long and 3-5 mm wide. Stems ascending, simple or sparsely branched, with lateral-intercalary or rarely terminal branching, with paraphyllia near the base of leaves and underleaves. Leaves contiguous to imbricate, obliquely inserted, succubous, somewhat antically secund, with the bases somewhat appendiculate, mostly 3-lobed up to 0.8-0.9 of the length; lobes subequal (ventral lobe shorter and narrower), ovate to lanceolate, convex, broadly recurved, with the ciliate margins (about 15-20 cilia on each margin); apex of 4-7 thick-walled, uniseriate elongate cells. Underleaves 2-lobed to 0.8-0.9 their length, mostly half the size of the leaves, lobes similar to leaf lobes. Cells more or less isodiametric to slightly elongated, with coarse nodose trigones mostly confluent, 18-25 × 15-20 µm, basal cells elongate to 30-40 × 15-20 µm. Cuticle smooth to finely papillose. Female bracts connate in ring, similar to leaves, but more ciliate to laciniate at margin. Perianth up to 1 cm long, deeply 6-plicate, with ciliate mouth. Seta with 10-12 cells in diameter. Capsule ellipsoidal. Spores papillose, 15-20 µm in diameter. Known only from boulders and as an epiphyte in mountain forests (Hagenia etc.) of the Rwenzoris and Mt Elgon. Also known from D.R. Congo, Ethiopia, Kenya, Rwanda and Tanzania.		a-f. Plicanthus giganteus: a, part of shoot, lateral view; b,c, leaves; d,e, underleaves; f, mid-leaf cells; Lisowski 3295 (BR) g-j. Plicanthus hirtellus: g,h, leaves; i,j, underleaves; O'Shea7401a (Herb. M.J. Wigginton). MALAWI.

Plicanthus hirtellus (F.Weber) R.M.Schuster, Nova Hedwigia 74: 492, 2002 (see figure above)

Plants in loose yellowish brown patches. Shoots rigid, 2-3 cm long and 1-2 mm wide. Stems procumbent with ascending tips, simple or sparsely lateral-intercalary branched, mostly without paraphyllia. Leaves contiguous to imbricate, obliquely inserted and succubous, in outline rounded-subquadrate, adaxially convex, unequally 3-lobed with dorsal lobe 2-3 × as long as ventral one, sinus acute to gibbous, descending up to 0.8-0.9 of the length of leaf; lobes acuminate, up to 10 cells long and 5 cells wide at base, with sharp marginal teeth or cilia (up to 5-7 on each margin); at least ventral lobe incurved. Underleaves large, deeply 2-lobed, about half the size of the leaves; lobes acuminate. Cells more or less isodiametric, marginal 14-16 µm, in the middle 15-20 × 10-15 µm, basal up to 30 µm elongated, trigones large, confluent. Cuticle verrucose. Oil-bodies 3-4 per cell, about 3-4 µm, finely granulate.

Known from humus on rocks, boulders and as an epiphyte in mountain forests of the Rwenzoris and from Bwindi National Park (Rukungiri). Also recorded in Africa from Bioko, Cameroon, Comoro Is., D.R. Congo, Ethiopia, Kenya, Mauritius, Madagascar, Malawi, Réunion, Rwanda, South Africa, Saõ Tomé, Seychelles, Sierra Leone, Tanzania and Zimbabwe. Widely distributed in eastern Asia (from Himalaya to Japan and Indonesia), also in northern Australia, Polynesia and Western Canada.

Tetralophozia (R.M.Schust.) Schljakov, Novit. Syst. Plant. non Vascul. 13: 227, 1976

Plants brownish to reddish brown. Shoots rigid, erect and filiform, mostly simple or very sparsely branched; branching lateral-intercalary, ventral-intercalary to terminal. Stem with 1-2-stratose cortex of strongly thick-walled cells. Leaves succubous, subtransversely inserted, hand-like, deeply (to 0.7-0.9 their length) 4-lobed, semiamplexicaul; lobes finger-like, abaxially sulcate, with basal part ciliate-laciniate. Underleaves 2-lobed, lobes leaflike. Cells small (mostly 10-20 µm) and firm-walled, with nearly absent trigones. Oil-bodies 2-4 per cell. Gemmae rare. Dioicous. Usually sterile or in unisexual populations. Androecia spicate, male bracts similar to leaves. Gynoecia terminal, female bracts similar to leaves, lobes more pointed and more toothed. Perianth emergent, deeply pluriplicate.

Tetralophozia cavallii (Gola) Vána, Trop. Bryol. 8: 102, 1993

*Plants* in loose tufts or scattered among other bryophytes, olive green to yellowish brown. Shoots rigid, filiform, 6-12 cm long. Stems erect to suberect, simple or sparsely branched; cortical cells in 2-3 rows thick-walled; paraphyllia absent (except in the female inflorescence). Leaves slightly to densely imbricate, nearly transversely inserted, not decurrent but semiamplexicaul, reniform, hand-shaped, deeply (to 0.7-0.9 their length) 4-lobed; lobes ovate-lanceolate, suberect to erect, subapiculate to acuminate, ending with some non-elongated cells, strongly channelled with reflexed margins and gibbous sinuses, with entire margins (at least in upper parts) except the basal and subbasal parts which are dentate by broad polymorphous laciniiform teeth; apical parts of cilia formed with strongly elongated cells. Apical cells in leaf lobes 10-15 µm, median cells 15-25 × 12-20 µm, basal only slightly elongated; thick-walled, with small and confluent or nearly absent trigones. Underleaves 2-lobed, large, of 0.7-0.8 the size of leaves, lobes similar, often falcate to acuminate, with cilia at base. Oil-bodies 2-6(-8) per cell, granular. Plants mostly sterile. Androecia intercalary, bracts 2-3 androus, slightly concave. Female bracts larger than leaves,		a, shoot; b,c, leaves; d, young leaf on stem, showing shape of insertion; e, underleaf; f, mid-leaf cells; g, cells of cilium; Porley M250a (Herb. M.G. Wigginton). MALAWI.

deeply 4(-5)-lobed, with dentate margins; at the bases of female bracts sometimes filiform bifid paraphyllia present. Perianth emergent, deeply 6-10-plicate, at least into middle part; perianth mouth ciliate.

Known only from rocks, boulders and as an epiphyte in mountain forests of the Rwenzoris, Mt Muhavura and Mt Elgon areas. Also known from D.R. Congo, Ethiopia, Kenya, Malawi, Rwanda, Tanzania and Zimbabwe.

Tritomaria Schiffn. ex Loeske, Hedwigia 49: 13, 1909

Plants in patches, mostly green. Shoots mostly 1-2 cm high, creeping to ascending with suberect apices. Stems with distinct thick-walled 1-3-stratose cortex, simple or rarely terminal to lateral-intercalary branched. Leaves subtransversely inserted, widely spreading, vertically oriented, concave, mostly asymmetrically 3-lobed, rarely 2- or 4-lobed, dorsal lobe abbreviated; lobes acute to obtuse, leaf margin entire, without cilia or teeth. Underleaves absent. Cells mostly thin-walled with distinct trigones; median and basal cells elongated. Cuticle mostly papillose. Gemmae elliptical to polygonal, mostly present on apices of leaf lobes. Dioicous plants. Androecia mostly intercalary, male bracts strongly saccate, 2-4-androus. Female bracts similar to leaves, sometimes slightly larger, 3-5-lobed, margins entire or irregularly dentate. Perianth obovoid to cylindrical, upper part 4-6-plicate, with contracted entire or dentate-ciliate mouth. Capsule wall 2-5-stratose.

Tritomaria exsecta (Schmidel) Schiffn. ex Loeske, Hedwigia 49: 13, 1909

Plants in loose to dense patches or scattered among other bryophytes, pale to deep green. Shoots mostly 1-2 cm long, flaccid. Stems prostrate, ascending to erect, mostly simple or only with subfloral innovations; cortical cells slightly smaller than the medullar ones. Rhizoids colourless. Leaves imbricate, suberect to spreading, canaliculate or concave, ovate to oval in outline, unequally (2-)3-lobed to 0.2-0.3 their length, dorsal lobe the smallest one; sinus shallow, obtuse to acute; lobes acute to acuminate. Cells thick-walled with minute trigones, apical 10-15 µm, median 10-20 × 8-15 µm, basal cells only slightly elongate. Cuticle striolate. Oil-bodies 2-7, in basal cells up to 15, spherical to ovoid, papillose. Gemmae in rust masses at apices of lobes of sterile plants, 2-celled, elliptical, 10-20 × 8-12 µm. Dioicous. Male plants with intercalary androecia, bracts saccate, 1-2-androus. Perianth cylindrical, mouth spinose-dentate. Spores 10-12 µm, papillose.

In Uganda collected only twice in montane forests: in the Rwenzoris, Nayabitaba Ridge, on very moist, flat-topped rock at 2620 m and on Mt Elgon, along Sasa trail, on decaying wood at 2700-3100 m.

a, shoot, dorsal view; b, shoot, lateral view; c-g, leaves; h, upper part of leaf showing shape of lobes; i, mid-leaf cells; Wood 1255 (E). Rwenzoris, Nayabitaba Ridge.

Also known from Ethiopia, Kenya, Malawi, Rwanda, South Africa and Tanzania. Widely distributed in the Holarctic, reaching from Japan through the Philippines to North Borneo; there is a very old report from Mexico.

Excluded species:
Andrewsianthus perigonialis (Hook.f. & Taylor) R.M.Schust.
The specimens from Mt Muhavura are misidentified: Hedberg 2223 p.p. (UPS) is a mixture of Anastrophyllum auritum (Lehm.) Steph. and Anastrophyllum minutum (Schreb.) R.M.Schust.; Hedberg 2073b (UPS) is Anastrophyllum auritum (Lehm.) Steph.