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Mosses and Liverworts of Uganda
JungermanniaceaeBy J. Vána (illustrations by M. C. Watling)Plants forming dense to loose patches or mats, rarely scattered among other bryophytes. Shoots small to robust, mostly 5-100 or more mm high and 0.1-10 mm wide, creeping, prostrate, ascending, suberect to erect. Stems mostly with weakly differentiated cortical cells, sometimes hyalodermis of larger cells present; branching not common, terminal or lateral-intercalary, very rare ventral-intercalary in form of stolons. Rhizoids scattered on the ventral face of stem, rarely originating from leaf base or leaf cells. Leaves mostly obliquely, rarely subtransversely inserted, succubous, alternate, dorsally extended to stem midline, not decurrent, slightly decurrent or long decurrent, mostly unlobed, rarely retuse to shallowly bilobed; leaf margins entire. Underleaves only in some genera; if present, small, lanceolate to bilobed. Leaf cells thin-walled, mostly collenchymatous, with distinct or bulging trigones, rarely parenchymatous, with missing trigones. Cuticle smooth to verruculose. Oil-bodies (1-) 2-9 (-12) per cell, mostly ovoid, granulose; rarely homogenous. Asexual reproduction only in some species, mostly by 1-2-celled gemmae; endogenous gemmae in one species (Jungermannia caespiticia). Dioicous or paroicous, exceptionally heteroicous plants. Androecia terminal, becoming intercalary or subgynoecial, spicate. Antheridia 1-3 (-6) per bract, ovoid, with 1-2-seriate stalk. Gynoecia terminal on main shoots, with 1 (-3) pairs of bracts. Perianth present (in some genera double), exserted or hidden by the bracts, sometimes reduced and partially replaced by perigynium. Seta with numerous cells rows. Capsule ovoid to spherical, 4-valved, with 2 (-5)-stratose wall; epidermal layer cells with nodular, inner layer cells with semiannular thickenings. Spores finely papillose; elaters mostly 2-spiral, rarely 1-or 3-4-spiral.
Jungermannia L., Spec. Plant. 1: 1131, 1753, emend. Dumort., Rec. Observ. Jungermanniacées, p. 16, 1835 Plants mostly forming green, brownish, reddish-brown to blackish patches or mats. Shoots small to robust, 5-250 mm long and 0.3-8 mm wide. Stems creeping, ascending to erect, with cortical cells hardly differentiated; branching not common, predominantly lateral-intercalary, rarely terminal. Rhizoids scattered, sometimes forming distinct bundle decurrent along the ventral side of stem. Leaves contiguous to imbricate, rarely remote, obliquely, rarely subtransversely inserted, sometimes decurrent, slightly concave to nearly plane, mostly ovate, cordate to rotundate, unlobed (exceptionally, in non-African species, retuse or slightly bilobed). Underleaves absent. Leaf cells small to large, mostly 20-30 µm, basal cells in some species more than 60 µm long, thin-walled, collenchymatous, rarely without trigones. Oil-bodies mostly 2-5 per cell. Dioicous, paroicous or rarely heteroicous. Androecia terminal, becoming intercalary, or subgynoecial, bracts in 2-20 pairs, ventricose at base, antheridia 1-3 (-6) per bract. Gynoecia terminal, sometimes with subgynoecial innovations. Bracts in 1 (-3) pairs, somewhat larger than vegetative leaves, bracteole only exceptional. Perianth cylindrical, ovate or obovate to fusiform; perigynium sometimes present. Seta mostly massive. Capsule oval to spherical, with 2 (-5)-stratose wall; epidermal cells mostly with nodular, inner wall with semiannular, sometimes incomplete semiannular thickenings. Spores mostly granular; elaters (1-) 2 (-3)-spiral. Key to fertile plants:
Key to sterile plants:
Jungermannia abyssinica Nees in Gottsche, Lindenb. et Nees, Syn. Hep., p. 93, 1844
In Uganda known only from the Rwenzori Mts, Mt Muhavura and Mt Elgon. On moist, exposed, steep rocks and gravely detritus, mainly in the alpine zone (3600-4000 m), though at only 2020 m by the Mahoma River (Rwenzori NP). An endemic of E. African mountains; known also from Ethiopia, Kenya and Tanzania.
-------------------- Jungermannia borgenii Gottsche in Pears., Christ. Vidensk.-Selsk. Forhandl. 1892: 11, 1893 (Fig. 2)
Mostly on soil or rock in the forest zone, ascending to about 2600 m in bamboo forest on Mt Elgon. Known only from Africa where it is the commonest species of the genus:Angola, Botswana, Cameroon, Congo (R. & D.R.), Kenya, Madagascar, Malawi, Mauritius, Nigeria, Réunion, Rwanda, Saõ Tomé, South Africa, Tanzania, Zambia, Zimbabwe.
--------------------- Jungermannia mildbraedii Steph. in Mildbraed, Wiss. Ergebn. deutsch. Zentral-Afrika Exped. 1907-1908, 2: 113, 1914 (Fig. 2) Plants in green, yellowish-green, olive brown to brownish-green patches. Shoots creeping, fertile shoots suberect, 5-30 mm high and 0.3-1.4 mm wide; branching lateral-intercalary, mostly in form of subfloral innovations. Stem with 1-3 rows of smaller (22-28 µm) cortical cells, subcortical cells 30-45 µm. Rhizoids relatively common, hyaline to brownish, diffuse, never in groups. Leaves obliquely inserted, mostly remote, ovate, ovoid, sometimes also cordate to rotundate, dorsally slightly decurrent. Marginal cells 14-18 µm, median ones 20-30 µm, basal elongated to 40 µm; trigones small. Cuticle mostly smooth. Oil-bodies 2-9 per cell, granular. Paroicous. Androecia subgynoecial, bracts in 1-3 pairs, sometimes separated from the gynoecium by 1-2 pairs of sterile leaves, antheridia (1-) 2 (-3) per bract. Gynoecia terminal, mostly with subfloral innovations, bracts mostly reniform, larger than the vegetative leaves. Perianth pyriform to fusiform, perianth mouth sometimes beaked; perigynium absent. Capsule spherical, with bistratose wall. Spores 16-18 µm; elaters 2-spiral. Collected only twice in Uganda, on wet rock in the alpine zone of the Rwenzori Mts: at 4400 m in a side valley on the S.E. slope of Mt Geesii by O. Hedberg in 1948, and at about 4850 m on S.E. side of Mt Stanley by E. Esterhuysen in 1956. Known only from East African mountains (Rwanda, Tanzania) and South Africa. See under Jungermannia borgenii, nos. f-j, for the illustration. -------------------- Jungermannia pumila With., An Arrang. British Plants, Ed. 3, 3: 883, 1796 (Fig. 3)
Collected only once in Uganda (by K. Wesche), from wet rock in the alpine zone of Mt Elgon. Also known from Lesotho, South Africa and Tanzania, widespread in the holarctic region and present on Java.
Jungermannia
renauldii. a, Shoot; b-d, leaves; e, mid-leaf cells; Herb. J.
Vána RÉUNION. -------------------- Jungermannia renauldii Steph., Bull. Soc. Roy. Bot. Belgique 30: 201, 1891 Plants in green to yellowish-green patches. Shoots creeping, 10-30 mm high and 0.8-2 mm wide; branching not common, mostly lateral-intercalary and as subfloral innovations. Stem with very weakly differentiated cortex of smaller cells. Rhizoids common, hyaline to yellowish, diffuse, never in groups. Leaves obliquely inserted, imbricate, ovate to oval, concave, slightly decurrent ventrally, long decurrent dorsally. Marginal leaf cells 22-30 µm, median ones 25-33 µm, basal cells elongated to 60 × 40 µm. Cuticle smooth or nearly so. Oil-bodies unknown. Dioicous. Male plants smaller than female ones. Antheridia terminal, becoming intercalary, bracts in 3-6 pairs, antheridia 1-2 per bract. Gynoecia terminal, sometimes with subfloral innovations, bracts larger than vegetative leaves. Perianth ellipsoid to fusiform, gradually contracted to the mouth; low perigynium always present. Capsule ovoid, with bistratose wall. Spores 14-15 µm, papillose; elaters 2-spiral. A single record (by K. A. Lye in 1969), from the edge of a swamp, Lake Kayanja, Masaka. Also known from Madagascar and Réunion. See under Jungermannia pumila, nos. f-j, for the illustration. -------------------- Jungermannia sphaerocarpa Hook., British Jungerm., tab. 74, 1815 (Fig. 4)
Known only in the vicinities of the Rwenzoris, Mt Muhavura and Mt Elgon, growing mostly on moist rocks, stones, earth banks and wet soil in the upper forest and alpine zones (3830-4000 m). Also recorded in Africa from Cameroon, D.R. Congo, Lesotho, Madagascar, Malawi, Réunion, Rwanda, South Africa, Tanzania and Zimbabwe. Widely distributed in the Holarctic and penetrating to the high mountains in the tropics in Borneo, Brazil, Columbia, Costa Rica, Ecuador, Mexico, New Guinea, Peru and Venezuela. Jungermannia sphaerocarpa. a, Shoot; b-e, leaves; f, basal to mid-leaf cells; De Sloover 18.920 (BR) RWANDA.
Nardia Gray, Nat. Arrang. Brit. Plants 1: 679 + 694, 1821 (“Nardius”), corr. Carrington, Trans. Proc. Not. Soc. Edinburgh 10: 309, 1870 (nom. cons.) Plants in green, yellowish-green, grayish-green, brownish to blackish patches or mats, sometimes tinged with red or deeply reddish. Shoots very small to large, 1-250 mm high and 0.12-10.0 mm wide. Stems creeping, prostrate to ascending, mostly without hyalodermis and with hardly defined 1-3 cortical layers of cells. Branching lateral-intercalary, terminal or rarely ventral-intercalary (N. compressa only). Rhizoids rare to frequent, scattered on the ventral face of stem. Leaves contiguous to remote, subtransversely to obliquely succubously inserted, broadly ovate, orbicular to reniform, entire, retuse to bilobed, sometimes shortly to longly decurrent, with entire margins. Underleaves small (in N. nuda only present on the stem apices), lanceolate to subulate, in some species connected with one leaf. Cells collenchymatous, small to medium sized, trigones small to nodulose. Cuticle mostly smooth. Oil-bodies 1-8 per cell, homogenous to granulose. No asexual reproduction. Dioicous or paroicous. Androecea subgynoecial or terminal, becoming intercalary, bracts hardly saccate, mostly emarginate or deeply bilobed, antheridia (1-) 2-3 per bract. Gynoecia terminal. Perianth mostly reduced (except N. nuda and some other taxa), replaced by fleshy stem-perigynium of Isotachis- or Nardia geoscyphus-type. Capsule ovoid to globose, with 2-stratose wall. Spores mostly small, 9-22 µm, elaters (1-) 2 (-4)-spiral. Nardia arnelliana Grolle, Bot. Magaz. Tokyo 77: 297, 1964 (Fig. 5) So far recorded from only a single locality in Uganda, on wet rock at about 4850 m on the S.E. side of Mt Stanley (collected by E. Esterhuysen in 1956), and from another locality in Réunion.
Reference Grolle R. 1964. Eine Nardia auf Afrika. Botanical Magazine, Tokyo 77: 297-299.
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Plants in
green, brownish-green to blackish-green patches. Shoots suberect
to erect, 10-30 mm high and 0.8-2 mm wide; branching relatively
rare, lateral-intercalary, mostly in form of subfloral innovations. Stem with
1-3 rows of cortical cells somewhat smaller (30-35 µm) as the
subcortical cells (40-45 µm). Rhizoids relatively
frequent, hyaline to brownish, originating from the ventral side of stem,
forming a distinct
group and ventrally decurrent along the stem; also rarely arising from
leaf cells. Leaves subtransversely inserted,
mostly imbricate, rotundate to reniform, weakly decurrent, with sheathing
base; leaf margin somewhat undulate. Marginal leaf
cells 20-30 µm, median cells 30-45 µm, basal
cells elongated to 90 × 45 µm; trigones small. Cuticle smooth. Oil-bodies unknown.
Paroicous. Androecia subgynoecial,
bracts in 1-4 pairs, saccate, antheridia 2-3 per bract. Gynoecia terminal,
bracts somewhat larger than the vegetative
leaves. Perianth ovoid to pyriform, contracted perianth mouth beaked; perigynium
low. Capsule spherical, with bistratose wall. Spores 25-27 µm, papillose;
elaters 2-spiral.
Plants in
green, yellowish-green to brownish patches, sometimes tinged with red.
Shoots creeping with ascending tips, 7-30 mm high and 1-2
mm wide; branching lateral-intercalary and ventral-intercalary; the latter
producing stolons. Stem with 1-2 rows of smaller (20-40 µm) cortical
cells, subcortical cells 35-45 µm. Rhizoids relatively common, hyaline
to brownish, diffuse, never forming a group. Ventral stolons sometimes present.
Leaves obliquely inserted, mostly imbricate, widely ovate to rotundate, slightly
decurrent. Marginal leaf cells 25-40 µm, median ones 30-45 µm,
basal cells elongated to 65 × 40 µm, trigones small to large.
Cuticle smooth. Oil-bodies 2-8 per cell, ovate, granular. Dioicous. Androecia terminal, rarely intercalar, bracts in 2-5 pairs, saccate, antheridia 1-3
per bract. Gynoecia terminal, sometimes with subfloral innovations; bracts
larger than vegetative leaves. Perianth ovoid, pyriform to shortly cylindrical,
suddenly contracted, perianth mouth mostly not beaked. Perigynium absent
or very low. Capsule spherical, with bistratose wall. Spores 14-20 µm,
papillose; elaters 2-spiral.
Plants in
green to olive-green, sometimes blackish patches, rarely intermixed among
other bryophytes. Shoots creeping with ascending
fertile stems, 5-30 mm high and 0.5-2 mm wide. Branching rare, mostly lateral-intercalary,
rarely terminal and exceptionally ventral-intercalary. Stem with 1-3 rows
of cortical cells, 30-35 µm across, the walls somewhat thickened, surrounding
smaller medullary cells 25-30 µm across. Rhizoids uncommon, scattered,
mostly hyaline. Leaves mostly distant, nearly flat, oval to broadly elliptical.
Leaf cells leptodermous, without trigones, marginal about 20-25 µm,
median ones 25-35 µm, basal sometimes elongated to 90 × 30 µm.
Cuticle verruculose. Oil-bodies 2-6 (-8) per cell, in basal cells sometimes
to 12 per cell, granulose. Paroicous. Androecia subgynoecial, bracts in 2-3
pairs, sometimes separated by sterile leaves from gynoecia, ovate, basally
saccate, mostly 1 antheridium per bract. Gynoecia terminal, bracts broadly
ovate. Perianth fusiform, pluriplicate, gradually narrowed towards crenulate
mouth; perigynium not developed. Capsule ovate, with bistratose wall. Spores 17-24 µm, papillose; elaters 2-spiral.
Plants in green to yellowish green flat mats,
sometimes tinged with red. Shoots very small,
to 10 mm high and 0.12-0.2 mm wide. Stems fragile,
creeping or ascending, with one row thin-walled cortical cells and thick-walled
intracortical cells. Branching lateral-intercalary,
rarely terminal. Leaves remote, transversely
inserted, concave, ovate to orbicular, not decurrent, bilobed to 0.2-0.3
their length; lobes obtuse to subacute. Underleaves subulate,
sometimes retuse on the tip. Leaf cells with
small trigones, about 10-15 µm, basal cells
not elongated. Cuticle smooth. Oil-bodies unknown. Paroicous. Androecia subgynoecial,
bracts in 2-3 pairs, much larger than vegetative leaves; antheridia 2
per bract. Gynoecia terminal, female bracts
larger than vegetative leaves. Perianth reduced,
conical, concealed by the bracts, partially replaced by perigynium of Isotachis-type. Sporophyte unknown.