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Mosses and Liverworts of Uganda

Fissidentaceae

By M. A. Bruggeman-Nannenga

This family includes only the genus Fissidens in Uganda, and the family description is therefore the same as that of the genus.


Fissidens Hedw., Sp. Musc., 1801

Plants minute to several cm long, sometimes with axillary nodules. Leaves arranged in 2 opposite rows, pinnate or flabelliform, each leaf consisting of a sheath composed of two vaginant laminae, an apical and dorsal lamina (Fig. 1b), leaves elimbate or partially to completely limbate, lanceolate, elliptical, oblong or lingulate; limbidium marginal or less often intralaminal; vaginant laminae slightly unequal to equal, infrequently strongly unequal with the smaller laminae ending on the costa; costa excurrent to ending far below leaf apex, infrequently greatly reduced to lacking; leaf cells smooth, unipapillose or pluripapillose, to 125 µm, but mostly less than 27 µm long, parenchymatous, infrequently prosenchymatous. Autoicous, synoicous, dioicous, often polyoicous. Perigonia axillary or terminal, perichaetia typically terminal. Seta mostly 1(–2) per perichaetium. Capsule erect, inclined or horizontal; peristome mostly are either bryoides–, scariosus–, zippelianus– (not in Uganda), similiretis– (not in Uganda) or taxifolius– (in Uganda in F. ovatus) types (Bruggeman–Nannenga & Berendsen, 1990). Operculum conic-rostrate. Spores finely papillose or smooth.

Cosmopolitan, aquatic, terrestrial, epilithic and epixylic, mostly in moist or wet places.


Notes on the key
In the descriptions and key below 'leaf cells' refers to cells of the mid dorsal lamina and 'leaves' to the middle leaves in pinnate plants, or to the one below the upper leaf pair below the upper leaf pair in flabelliform plants.

Expressions of variable species from outside Uganda may key out incorrectly with the key below.


1. Leaves with distinct costa 2
  Most leaves ± without costa, infrequently perichaetial leaves with a barely visible costa 3
     
2. Leaves lacking dorsal and apical laminae F. gladiolus
  Dorsal and apical lamina present in all well-developed leaves 5
     
3. Leaves elimbate, marginal cells mostly smaller than inner leaf cells, perichaetial leaves often with inconspicuous costa F. usambaricus
  Leaves with border of long and narrow cells (limbidium), perichaetial leaves without costa 4
     
4. Limbidium ill-defined, limbidial cells 4.5–9 µm wide; often with clusters of gemmae on leaves F. metzgeria
  Limbidium distinct, limbidial cells ± 1 µm wide, leaves without gemmae F. enervis
     
5. Leaf cells 27 µm or more long F. flaccidus
  Leaf cells less than 27 µm, mostly not over 22.5 µm long 6
     
6. Costa over its whole length obscured by short, wide cells F. cryptoneuron
  Costa distinct, not obscured by short, wide cells 7
     
7. Dorsal and apical laminae with a dark border of short wide cells often mixed with a few elongated ones F. porrectus
  Dorsal and apical lamina without dark border of short wide cells 8
     
8. Smaller vaginant laminae rounded and free above, ending on or near the costa; leaf apex rounded with costa ending 7–9 cells below F. asplenioides
  Smaller vaginant laminae not rounded and free above, mostly ending close to the leaf margin, infrequently on or near the costa; leaf apex and costa various 9
9. Leaves narrow, 6–7 times as long as wide; limbidium restricted to the vaginant laminae, or absent; costa ending 15–26 cells below leaf apex F. palmifolius var. semilimbatus
  Leaves wider, or, when 6 or more times as long as wide, limbate on all laminae; costa percurrent, excurrent or ending 1–4(–11) below the leaf apex 10
     
10. Large axillary nodules present (Fig. 15a, h), leaves elimbate F. ovatus
  Axillary nodules absent or small (Fig. 6h), leaves limbate or not 11
     
11. Leaves limbate on vaginant, apical and dorsal laminae 12
  Leaves elimbate, or if limbate, limbidium restricted to the vaginant laminae 25
     
12. Leaf cells smooth, flat or convex 13
  Leaf cells papillose or when smooth conical 21
     
13. Female stems mostly shorter with fewer, but larger leaves than infertile plants 14
  No marked difference between stems and leaves of female and infertile plants 16
     
14. Vaginant laminae strongly unequal, smaller one ending on the costa F. beckettii
  Vaginant laminae not or slightly unequal, smaller vaginant lamina ending at or near the margin 15
     
15. Limbidium and costa mostly confluent at leaf apex, limbidium marginal; costa percurrent to long excurrent F. curvatus
  Limbidium ending below leaf apex, intralaminal in lower part of vaginant lamina; costa ending 1–2 cells below leaf apex F. bogosicus
     
16. Capsule with ± 32 files of exothecial cells; scariosus-type peristome; plants mostly flabelliform; lower cells of vaginant laminae pellucid, to 37.5 µm long

F. zollingeri

  Capsule with 40 or more files of exothecial cells; bryoides-type peristome; plants mostly pinnate; lower cells of vaginant laminae not pellucid, mostly shorter than 37.5 µm 17
     
17. Lower 1/5 of dorsal laminae composed of long, narrow cells (indistinguishable from limbidium or costa cells); leaves narrowly elliptic-lanceolate F. robynsianus
  Lower 1/5 of dorsal laminae clearly distinct from costa and limbidium tissue; leaf shape variable 18
     
18. Leaf cells convex, bulging, small , 6–7.5 x 3–6 µm 19
  Leaf cells flat or lowly convex, not or slightly bulging, mostly larger 20
     
19. Vaginant laminae ± equal with limbidia distally confluent, mostly marginal throughout F. schmidii
  Vaginant laminae unequal with limbidia distally not confluent, intralaminal below F. megalotis subsp. helictocaulos
     
20. Limbidium ending below leaf apex; aquatic, often inundated F. leucocinctus
  Limbidium nearly or completely reaching the leaf apex; typically in dryer habitats F. androgynus
     
21. Leaf cells mammillose or unipapillose 22
  Leaf cells pluripapillose 23
     
22. Leaf cells 7.5–10.5 µm long F. angustifolius
  Leaf cells 3–4.5 µm long F. unipapillosus
     
23. Leaf cells large, 10.5–15 x 7.5–10.5 µm, clear, papillae minute F. curvatus
  Leaf cells smaller, obscure, papillae larger 24
     
24. Leaves stiff and narrow, 7–8 times as long as wide, limbidium of vaginant laminae 10 or more cells wide F. glaucissimus
  Leaves undulate, broader, 4–5 times as long as wide, limbidium of vaginant laminae narrower less than 6 cells wide F. weirii
     
25. Vaginant laminae of all leaves elimbate 26
  Vaginant laminae of some or all leaves partly or completely limbate 27
     
26. Leaf cells smooth, often guttulate F. pseudoeenii
  Leaf cells pluripapillose F. sciophyllus
     
27. Limbidium present on the entire length of the vaginant laminae of all leaves, often extending onto the adjacent part of the apical lamina 28
  Limbidium shorter, not necessarily present on all leaves, often restricted to upper leaves of perichaetial plants 29
     
28. Leaf cells mammillose with 1(–2) high and sharp papillae F. submarginatus
  Leaf cells ± flat, with 2–6 low, blunt papillae F. intromarginatus
     
29. Leaf cells smooth, flat or slightly convex, often guttulate F. pellucidus
  Leaf cells papillose or mammillose 30
     
30. Leaf cells unipapillose or mammillose 31
  Leaf cells pluripapillose 32
     
31. Peristome teeth straight, not or irregularly divided; vaginant laminae of female stems at base rounded and wider than the stem F. reflexus
  Peristome teeth curved, divided into 2 long, twisted, ± equal prongs; vaginant laminae of female stems narrower than the stem below F. ramulosus
     
32. Limbidia restricted to some or all leaves of fertile plants 33
  Limbidium present on most or all leaves of both fertile and infertile plants 34
     
33. Leaves ending in 1–5 pointed clear cells; costa percurrent, or ending 2-4(-8) below leaf apex F. sciophyllus
  Leaves not ending in a clear apical cell; costa ending below apex F. pallidinervis
     
34. Limbidium on sterile stem leaves extending less than ¼ of way up vaginant laminae, marginal to weakly intralaminal by one row of cells F. marthae
  Limbidium on sterile stem leaves reaching 1/2 or more the length of the vaginant laminae 35
     
35. Limbidia marginal F. cuynetii
  Limbidia intralaminal, bordered by 1–3 rows of cells F. lachmanii

Notes on the species descriptions

Unless stated otherwise all species below have pinnately arranged leaves, and lack persistent protonemata, gemmae, tubers and axillary nodules (i.e. axillary cells can be differentiated or not, but they do not protrude).

The bryoides-type peristome is amongst other features characterized by wide-spaced horizontal ridges on the basal part of the peristome teeth (Fig. 4c) and is found in combination with 40 or more files of exothecial cells.

The scariosus-type peristome is among other features characterized by many horizontal ridges on the basal part of the peristome teeth (Fig. 12g) and is found in combination with ± 32 files of exothecial cells.

 

Fissidens androgynus Bruch in C.Krauss, Flora 29: 134, 1846 (Fig. 1)

 

 

Plants 6.5 x 3 mm. Leaves limbate, lanceolate to oblong-lanceolate, apex acute, often mucronate, 1.2–1.9 x 0.35–0.45 mm, 3-4 times as long as wide; limbidium marginal, in lower part of vaginant laminae often weakly intralaminal, reaching insertion or ending just above, in leaf apex often confluent with costa; vaginant lamina 3/5 the leaf length, slightly unequal to equal; dorsal lamina tapering to rounded at base, reaching the insertion, decurrent or not; costa often excurrent; cells 7.5–13.5 x 4.5–10.5 µm, smooth, ± flat. Perigonia, perichaetia and synoecia terminal on plants or branches; perichaetial leaves 1.5–2.5 mm long. Seta 6–8 mm long. Capsule ± erect, 1.1 x 0.5 mm, exothecial cells in ± 52 files. Peristome bryoides-type, teeth 52.5 µm wide at base. Operculum 0.4–0.5 mm long. Spores 13.5–15 µm diameter.

F. androgynus is recognized by its limbate leaves and smooth leaf cells. This species could be confused with F. schmidii which, however, has smaller, 6–7.5 x 3–6 µm, bulging leaf cells.

In Uganda known from: Kabale, Kapchorwa, Kabarole, Kigezi, Kisoro, Mbale, and Rukungiri Districts. Further African distribution: Angola, Cameroon, Cape Verde Islands, Central African Republic, D.R. Congo, Ethiopia, Equatorial Guinea (Bioko), Gabon, Ghana, Guinea, Ivory Coast, Kenya, Lesotho, Malawi, Mascarene islands, Nigeria, South Africa, Rwanda, Tanzania, and Zimbabwe.

Mostly on dead or living wood or rock, also frequently on soil; in shady, humid places, rarely submerged, 770–3960 m.


Figure 1. Fissidens androgynus: (a) upper part of plant; (b) leaf; (c) leaf apex; (d) insertion of vaginant lamina; (e) margin mid-dorsal lamina. Porley U9056A (E).

 


 Fissidens angustifolius Sull., Proc. Amer. Acad. Arts 5: 275. 1861 (Fig. 2)

Plants ± 3.5 x 3 mm, flabelliform, with hyaline, axillary nodules. Leaves limbate, narrowly elliptical, apex acute, in upper leaves cuspidate, middle leaves 1.9–2 x 0.35–0.5 mm, 4–5.5 times as long as wide. Limbidium marginal, on all laminae of upper and middle leaves, ending above insertion, reaching the leaf apex in upper leaves. Vaginant laminae 1/2 the leaf length, ± equal. Dorsal lamina tapering below, reaching the insertion. Costa percurrent to excurrent. Leaf cells 7.5–10.5 x 4.5–7.5 µm, mammillose to unipapillose. Perigonia bud shaped in leaf axils; synoecia and perichaetia terminal. Perichaetial leaves 2.5 mm long. Seta 4 mm long, capsule erect, 0.7 x 0.3 mm. Peristome scariosus-type, teeth 37.5 µm wide at base. Operculum 0.5 mm long. Spores 9–10.5 µm diameter.

This species is characterized by flabelliform plants, limbate leaves, distinct hyaline axillary nodules and mammillose to unipapillose leaf cells. It resembles F. zollingeri which, however, has flat to slightly convex leaf cells.

In Uganda rare, only known from Bushenyi, Kalinzu FR. Also known from Central African Republic, Gabon, Nigeria, and Togo. Outside Africa it is known from Mexico, South and Central America, and Polynesia.

On soil, bark, rotten wood and rock, to 1960 m.


 

Fissidens asplenioides Hedw., Sp. Musc. Frond.: 156, 1801 (Fig. 3)

Plants 3–30 x 1.5–4 mm, axillary nodules absent to well differentiated. Leaves elimbate, oblong-lanceolate, apex widely acute to obtuse, rounded, 1–2.5 x 0.25–0.6 mm, 4–5 times as long as wide. Vaginant laminae 3/5 the leaf length, strongly unequal, the minor one rounded and free distally, ending on or near the costa. Dorsal lamina mostly ending far above insertion. Costa ending 7–9 cells below leaf apex. Leaf cells 4.5–7.5 x 3–6 µm, smooth, strongly bulging, basal marginal cells of vaginant laminae linear. Dioicous. Perigonia and perichaetia terminal on plants or on branches from upper part of stem. Perichaetial leaves ± 4 mm long. Sporophytes rare, not known from Uganda.

In the dry condition this species is recognized by leaves strongly inrolled from tips. When wet it is recognized by its elimbate, oblong-lanceolate leaves with rounded apex, a costa that ends 7–9 cells below the leaf apex and strongly unequal vaginant laminae of which the smaller one is round and free above.

In Uganda known from: Kabarole, Kapchorwa, Kigezi, Kisoro, Mbale, Mengo and Rukungiri Districts. Widespread in Africa. Cameroon, Comores, Democratic Republic of the Congo, Ethiopia, Kenya, Macaronesia, Madagascar, Malawi, Nigeria, Guinea, Réunion, Rwanda, South Africa, Tanzania, Zambia and Zimbabwe Also found in the Americas, Australasia and the Pacific.

On wet and moist rock, and soil, often near streams, to 3350 m.


 

Fissidens beckettii Mitt., J. Linn. Soc. Bot. 13: 225, 1873 (Fig. 4a-e)

Plants dimorphic, infertile stems 3–5 x 1 mm; female stems 4–4.5 x 1.7–2 mm. Leaves limbate, ovate-lanceolate to elliptic-lanceolate, apex acute and long acuminate, 0.55–0.6 x 0.2 mm, 3 times as long as wide (leaves of female stems 1.1 x 0.33 mm; 4 times as long as wide). Limbidium marginal, confluent with the costa at apex of minor vaginant lamina, ending below or becoming indistinct in the leaf apex, reaching the insertion of the dorsal lamina or not. Vaginant laminae 1/2 the leaf length, unequal, smaller one ending on the costa. Dorsal lamina more or less rounded at base, reaching the insertion. Costa long excurrent. Leaf cells 10.5–15 x 7.5–9 µm, smooth and flat. Perichaetia terminal. Perichaetial leaves 1.7 mm long. Seta 4 mm long; capsule inclined, 0.4 x 0.3 mm, ± 42 columns of exothecial cells. Peristome bryoides-type, teeth 30 µm wide at base. Spores 13.5–15 µm. Perigonia and operculum not seen in material from Uganda.

This species is characterized by dimorphic plants, limbate, acuminate to narrowly acute leaves, unequal vaginant laminae with the smaller ones ending on or near the costa, excurrent costa and smooth leaf cells.

In Uganda known only from the Arua District. Widespread, but uncommon in Africa: Central African Republic, Republic of South Africa, Tanzania, and Zimbabwe. Also known from Asia, Mexico and the Caribbean.

On soil, infrequently on bark, moist, shady places, to 2000 m.


 

Fissidens bogosicus Müll.Hal. ex Vent., Nuovo Giorn. Bot. Ital. 4: 10, 1872 (Fig. 4f-k)

Plants dimorphic, infertile plants rare, pinnate, 5 x 1.5 mm; fertile plants flabelliform, 3.4 x 1.2 mm. Leaves limbate, elliptic-lanceolate, apex acute, acuminate, 1–1.3 x 0.3–0.35 mm, 3–3.5 times as long as wide. Limbidium marginal, intralaminal in lower part of vaginant laminae ending distinctly below leaf apex and above the insertion of dorsal lamina. Vaginant laminae half as long as the leaf, slightly unequal. Dorsal lamina reaching insertion, not decurrent. Costa ending 1–2 cells below leaf apex. Leaf cells 7.5–13.5 x 6–9 µm, smooth, flat to slightly convex. Perigonia terminal on short branches, some antheridia naked in upper leaf axils; perichaetia and synoecia terminal. Perichaetial leaves 1.2–1.4 mm long. Peristome bryoides-type (no sporophytes seen in Uganda material).

This variable species is characterized by dimorphic plants, wide, elliptic-lanceolate leaves, large leaf cells and bryoides-type peristomes. Leaves can be completely limbate as in the Ugandan material, partly limbate or elimbate; leaf cells are smooth in some collections (e.g. the Uganda specimen), but inconspicuously pluripapillose in others.

In Uganda rare, known only from Queen Elizabeth National Park, Bushenyi District. Endemic to Africa: Botswana, Cape Verde Islands, Democratic Republic of the Congo, Eritrea, Ethiopia, Kenya, Lesotho, Madagascar, Malawi, South Africa, Tanzania, Zambia, and Zimbabwe.

On rocks, soil, roots and bark, humid to dry places, to 2460 m.


 

Fissidens cryptoneuron P.de la Varde, Rev. Bryol. Lichénol. 5: 208, 1933 (Fig. 5a-e)

Plants 2–4 x 0.7–1.2 mm. Leaves elimbate to partially limbate, oblong-lanceolate, apex widely acute to obtuse, often ending in a clear top cell, 0.35–0.4 x 0.1–0.2 mm, 3–3.5 times as long as wide. Limbidium marginal, restricted to ¾ or less of the vaginant laminae of upper leaves of female stems, interrupted and weak. Vaginant laminae ±1/2 the leaf length, gaping, unequal. Dorsal lamina tapering below, often ending above insertion. Costa over its whole length covered by chlorophyllose cells, ending 10–19 cells below leaf apex. Leaf cells obscure, 6–7.5 x 4.5–7.5 µm, with 2–5 papillae. Perigonia bud-shaped and axillary or terminal on stems or branches; perichaetia terminal. Perichaetial leaves 0.9–1.1 mm long. Seta 1.5–3.0 mm long; capsule erect, 0.5–0.7 x 0.2–0.35 mm, ± 32 files of exothecial cells. Peristome teeth 33 µm wide at base. Operculum 0.25–0.4 mm long. Spores 19.5–21 µm.

This small species is unmistakable. It is at once recognized by short costae that are obscured by chlorophyllose cells over the whole length and pluripapillose leaf cells.

In Uganda rare: Olwal Forest Reserve (Gulu District) and Arua District. Widespread, but uncommon: Central African Republic, Democratic Republic of the Congo, Guinea, Kenya, Madagascar, Tanzania, and Zambia. Also known from Brazil.

On bark and stone, 550–2400 m.


 

Fissidens curvatus Hornsch., Linnaea 15: 148, 1841 (Fig. 5f-j)

Plants dimorphic, infertile plants pinnate, 6 x1.7 mm, fertile plants flabelliform, 2.5–3.5 x 1.7–2 mm. Leaves limbate, elliptic-lanceolate, apex acute to acute-acuminate, cuspidate in female plants, 0.7–1.2 x 0.2–0.4 mm, 3–4 times as long as wide. Limbidium marginal, extending from the base of vaginant and dorsal laminae to the leaf apex. Vaginant laminae 3/5 the leaf length, slightly unequal to equal. Dorsal lamina tapering below, reaching the insertion, decurrent or not. Costa percurrent to excurrent. Leaf cells 10.5–15 x 7.5–10.5 µm, smooth or finely pluripapillose. Perigonia terminal on dwarf male plants at base of female plants or bud-shaped in leaf axils; perichaetia terminal on plants or branches. Perichaetial leaves 1.5 mm long. Seta 5–11 mm long; capsule erect to slightly inclined, 0.9–1.1 x 0.2–0.5 mm, ± 62 columns of exothecial cells. Peristome bryoides-type, teeth 40.5 µm wide at base. Operculum not seen. Spores ±15 µm long.

This species is characterized by narrow, lanceolate, limbate leaves with acute to cuspidate tips, dimorphic stems and large leaf cells that are smooth in some collections and smooth looking, yet delicately pluripapillose in others. The specimen from Rukungiri District is a poorly limbate form.

In one collection radiating, strongly birefringent, colourless, multicellular, pointed outgrowths (gemmae?) were found on some rhizoids.

In Uganda rare: Arua and Rukungiri Districts, and Mount Elgon (although this latter collection may be from the Kenyan side of the border). Widespread in Africa: Canary Islands, Ethiopia, Equatorial Guinea (Bioko), Lesotho, Madagascar, Madeira, Malawi, South Africa, Swaziland, Tanzania, and Zimbabwe. Further known from Europe, Australia, New Zealand, New Caledonia, Asia, South and North America.


On soil, rock, rarely on wood, to 3600 m.


 

Fissidens cuynetii Bizot ex Pócs, Fol. Hist.–Nat. Mus. Matr. 4: 29, 1977 (Fig. 6)

Plants 4–9 x 2 mm. Leaves ± semilimbate, oblong-lanceolate, widely acute to obtuse, often rounded-apiculate, 1.4 x 0.37–0.4 mm, ± 3.5 times as long as wide. Limbidium marginal, extending 4/5 the length of the vaginant laminae of all or most leaves. Vaginant laminae ¾ the leaf length, in upper leaves often with reflexed margins, slightly unequal. Dorsal lamina somewhat rounded and undulate at base. Costa percurrent to ending 2–4 cells below leaf apex. Leaf cells 4.5–7.5 x 4.5–7.5 µm, with 3–5 papillae. Perichaetia terminal. Perichaetial leaves 1.5 mm long. Perigonia and sporophyte unknown.

This species resembles F. intramarginatus in its pluripapillose leaf cells and semilimbate vaginant laminae. In F. cuynetii, however, limbidia are restricted to 4/5 of the vaginant laminae, whereas those of F. intramarginatus extend the whole length of the vaginant laminae and often a little onto the apical laminae.

In Uganda: near Motozho (Bushenyi), from near Nakuwa Lake (Kamuli), and Musoma, Damba Island (Mukono). A rare African species known only from Uganda and Rwanda.

On soil and rock, 1300–1700 m.


 

Fissidens enervis Sim, Trans. Roy. Soc. South Africa 15: 187, 1926 (Fig. 7a-c)

Plants 4.5–7 x 1.2–2 mm. Leaves limbate, elliptical, apex acute, 1.2–2.1 x 0.5–0.6 mm, 2.5–4 times as long as wide. Limbidium marginal, 4.5–15 µm wide, ± reaching the leaf apex and the insertion of the dorsal lamina. Vaginant laminae half or less the leaf length, equal or slightly unequal. Dorsal lamina tapering towards the insertion, reaching the insertion, not decurrent. Costa ± absent. Leaf cells large, 45–67.5 x 22.5–37.5 µm, smooth. Perichaetia terminal. Perichaetial leaves 1.5–1.7 mm long. Perigonia and sporophytes not seen in Uganda material.

This species is characterized by leaves without costae, distinct limbidia of linear cells and large leaf cells. It could be confused with F. metzgeria which, however, has wider, ill-defined limbidia of wider cells.

In Uganda uncommon: Arua, Kabarole, and Kisoro Districts. Further distribution: Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko), Kenya, Malawi, South Africa, Tanzania, Zambia, and Zimbabwe. Endemic to Africa.

On soil, infrequently on rocks or roots, on road-banks and of gullies, 610–2510 m.


 

Fissidens flaccidus Mitt., Trans. Linn. Soc. London 23: 56, 1860 (Fig. 8)

Plants 7–6 x 1.5–3.5 mm. Leaves limbate, oblanceolate, apex acute, ending in a pointed, often coloured cell, 1.5–3 x 0.3–0.6 mm, 4–5 times as long as wide. Limbidium marginal, reaching the insertion, confluent at the leaf apex. Vaginant laminae ½–3/5 the leaf length, equal. Dorsal lamina tapering below, reaching insertion, not decurrent. Costa ending 10–16 cells below leaf apex. Leaf cells large, 30–61.5 x 10.5–18 µm, flat, slightly convex or lowly conical. Perichaetia terminal. Perichaetial leaves 3.1 mm long. Perigonia and sporophytes unknown from Uganda.

This species is easily recognized by its limbate leaves, large leaf cells and short costae. Axillary bundles of rhizoids ending in filamentous gemmae are occasionally found.

In Uganda known from: Arua, Kabarole, and Mbale Districts. Widespread in Africa: Cape Verde Islands, Central African Republic, Ivory Coast, Nigeria, Rodriguez, Seychelles, Sierra Leone and Zimbabwe. Also in the neotropics.

On soil, rock or wood. Frequently along streams and waterfalls. In Africa to 1585 m, in the neotropics to 3350 m.


 

Fissidens gladiolus Mitt., Trans. Linn. Soc. London 23: 55, t. 6, f. 16, 1–3, 1860 [1862] (Fig. 9)

Fertile plants (infertile plants not seen) 1–2.5 x 1–1.5 mm, protonemata persistent. Leaves elimbate, linear-lanceolate, slightly unequal, subulate, perichaetial leaves 1–2.5 x 0.2 mm, 8-10 times as long as wide. Dorsal and apical lamina not developed. Costa long excurrent. Leaf cells linear-rectangular to linear-hexagonal, 51–105 x 4.5–9 µm, smooth. Perigonia terminal on dwarf plants at base of female plants; perichaetia terminal. Seta 6 mm long; capsule strongly inclined to curved, 0.8 x 0.4 mm, ± 56 columns of exothecial cells. Peristome bryoides-type, teeth 45–46.5 µm wide at base. Operculum 0.5 mm long. Spores 10.5–12 µm long.

This small species is probably often not recognized as Fissidens. The plants are too short to be conspicuously distichous, and the dorsal and apical laminae are often lacking as in the Uganda collections. However, the bryoides-type peristome eliminates all doubts about its place being in the Fissidentaceae. Once it is recognized as Fissidens, identification is easy. The species is characterized by persistent protonemata, subulate leaves often lacking a dorsal and apical lamina, linear leaf cells and bryoides-type peristomes. The vaginant laminae are elimbate or have an ill-defined limbidium, the dorsal and apical laminae, when present, are limbate. A similar species occurs in Australia.

Uganda: Kibale NP (Kabarole District) and Budongo Forest (Masindi). This uncommon African species is also known from the Central African Republic, Gabon, Nigeria, South Africa and Tanzania.

On (upturned) soil, clay, once on a termite mound, 950–2000 m.


 

Fissidens glaucissimus Welw. & Duby in Duby, Mém. Soc. Phys. Genève: 9, 1871 [1870] (Fig. 10)

Plants 5–10 x 2–3 mm. Leaves limbate, linear lanceolate, 1.9–2.1 x 0.28 mm, 7–8 times as long as wide. Limbidium marginal, ± reaching leaf apex, ending above insertion of dorsal lamina. Vaginant laminae ¾ the leaf length, slightly unequal. Dorsal lamina reaching the insertion, not decurrent. Costa percurrent to long excurrent. Leaf cells obscure, 6 X 4.5–6 µm, with to 8 marginal papillae. Perigonia terminal on 0.5–2 mm long plants at foot of female plants; perichaetia terminal. Perichaetial leaves 2.3 mm long. Seta 3–7.5 mm long, 1–2 per perichaetium; capsule inclined to horizontal, 0.8–1.2 x 0.45–0.6 mm, 32(–42) files of exothecial cells. Peristome scariosus-type, teeth 32–49.5 µm wide at base. Operculum 0.6 mm long. Spores 7.5–12 µm long.

This species is unmistak
able by its narrow, limbate leaves, pluripapillose, dark leaf cells and limbidia that are 9–14 cells wide on the vaginant laminae.

In Uganda known from: Kampala, Kitgum, Masindi, Mukono and Rukungiri Districts. Widespread in Central Africa: Angola, Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, Guinea, Ivory Coast, Kenya, Nigeria, Príncipe, São Tomé, Tanzania, and Togo. Endemic to Africa.

On soil and rock, infrequently on termite mounds or decaying wood, in humid places, to 2485 m.


 

Fissidens intramarginatus (Hampe) A.Jaeger, Enum. Fissident. 14, 1869 (Fig. 11)

Plants 6.5–9 x 1.9–3.6 mm. Leaves semilimbate, lanceolate to elliptico-lanceolate, apex acute, 1.8–2.1 x 0.4–0.6 mm, 3.5–5 times as long as wide. Limbidium marginal, on vaginant laminae of all leaves, extending the length of the vaginant laminae and often slightly onto the apical lamina, infrequently a few limbidial cells in the middle of the dorsal laminae. Vaginant laminae 3/5 the leaf, slightly unequal, margins often reflexed. Dorsal lamina rounded at base, reaching insertion, not decurrent. Costa percurrent to excurrent. Leaf cells 6–10.5 x 3–4.5 µm, with 2–7 papillae. Perigonia terminal on short plants at base of female plants; perichaetia terminal, a few archegonia axillary, naked. Perichaetial leaves 2.2 mm long. Seta 4.5 mm long; capsule erect to inclined, 0.9 X 0.3 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 39 µm wide at base. Operculum 0.7 mm long. Spores 7.5–10.5 µm long.

This variable species is characterized by limbidia extending the whole length of the vaginant laminae and often shortly onto the apical lamina and pluripapillose leaf cells. Fissidens submarginatus is also semilimbate, but has unipapillose, though often mixed with bipapillose ones, leaf cells. Papillae of F. submarginatus are typically high and sharp, whereas those of F. intramarginatus are low and blunt.
Several collections from Uganda have axillary clusters of filamentous gemmae.

In Uganda known from: Arua, Bushenyi, Kabale, Kalanagala, Kabarole, Luwero, Masaka, Masindi, Mpigi, Mubende, Mukono and Rukungiri Districts. Widespread in Africa: Benin, Cameroon, Central African republic, Democratic Republic of the Congo, Gabon, Equatorial Guinea (Bioko), Ghana, Guinea, Ivory Coast, Kenya, Liberia, Madagascar. Malawi, Nigeria, Réunion, Rwanda, Senegal, Sierra Leone, Sudan, Tanzania, and Togo. Also widespread in the neotropics.

Mostly on soil or rock, also decaying wood, termite mounds, roots and tree-ferns, to 3050 m.


 

Fissidens lachmannii Bizot, Rev. Bryol. Lichénol 40 (2): 134, 1974 (Fig. 12)

Plants 5–8 x 1.3–2.7 mm. Leaves ± semilimbate, oblong-lanceolate, acute, 0.95–1.3 x 0.27–0.4 mm, 3–3.5 times as long as wide. Limbidium intralaminal by 1–3 rows of cells, extending 3/5–5/7 the length of vaginant laminae of all or most leaves, often interrupted. Vaginant laminae about ½ the leaf length, slightly unequal. Dorsal lamina rounded-truncate at base, reaching insertion, not decurrent. Costa ending 1–3 cells below apex. Leaf cells 3–6 x 3–4.5 µm, with 4–7 marginal papillae. Perigonia bud-shaped, axillary; perichaetia terminal on branches from upper part of female stems. Perichaetial leaves 1.3–1.6 mm long. Seta 2 mm long; capsule slightly oblique to erect, 0.6–8 x 0.35 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 40.5–45 µm wide at base. Operculum not seen. Spores 9–13.5 µm long.

This species is characterized by short, intralaminal limbidia on all or most leaves and pluripapillose leaf cells.

In Uganda known from Bushenyi and Kabarole Districts. Elsewhere the species has been reported from Cameroon and Ivory Coast. Endemic to Africa.

On bark of tree trunks, roots and branches, to 1350 m.


 

Fissidens leucocinctus Hampe, Linnaea 38: 222, 1874 (Fig. 13a-e)

Plants 9–15 x 2–2.5 mm. Leaves limbate, broadly elliptical, elliptic-lanceolate or ovate, apex broadly acute, 1.2–2 x 0.35–0.7 mm, 3–4 times as long as wide. Limbidium marginal, frequently intralaminal in basal part of vaginant laminae, ending below leaf apex and often above the insertion of dorsal lamina. Vaginant laminae ½–3/5 the leaf length, equal. Dorsal lamina tapering below, reaching the insertion, not to slightly decurrent. Costa ending 2–3 cells below leaf apex. Leaf cells 6–9 x 4.5–7.5 µm, smooth, convex or flat, uni- to bistratose. Perichaetia terminal. Perichaetial leaves 2.2 mm long. Perigonia and sporophytes not known from Uganda.

This aquatic species is characterized by broad, limbate leaves that are strongly narrowed below and at apex, and by small, smooth, often slightly convex leaf cells. It can be confused with F. schmidii which, however, has narrower, lanceolate leaves, bulging leaf cells and is not aquatic.

In Uganda known from Kabarole, Kapchorwa and Rukungiri Districts. Further distribution: Cameroon, Democratic Republic of the Congo, Gabon, Kenya, Madagascar, Réunion, Tanzania, and Zimbabwe. Endemic to Africa.

On rock and wood in river beds and near waterfalls, often inundated, 480–2100 m.


 

Fissidens marthae Cardot, Rev. Bryol. 35: 64, 1908 (Fig. 13f-j)

Plants 3–5 x 1.7 mm. Leaves with short limbidia, oblong-lanceolate to oblanceolate, apex widely acute with indistinct, blunt apiculus, 1.1–1.3 x 0.3–0.35 mm, 3.5 times as long as wide. Limbidium marginal to weakly intralaminal by one row of cells, restricted to the basal 1/5 (2/3 in perichaetial leaves) of the vaginant laminae of all or most leaves. Vaginant laminae 3/5 the leaf length, slightly unequal. Dorsal lamina slightly rounded below, reaching the insertion, not decurrent. Costa ending 2–3 cells below leaf apex. Leaf cells 3–7.5 x 3–6 µm, with 1–4 (–7) low papillae, often a few cells protruding from surface of the lamina and/or bistratose. Perigonia terminal on dwarf plants; perichaetia terminal, infrequently a few naked, solitary, axillary archegonia. Perichaetial leaves 1.7 mm long. Seta 2 mm long; capsule ± erect, 0.7 x 0.5 mm, ± 38 files of exothecial cells. Peristome scariosus-type, teeth 45 µm wide at base. Operculum 0.6.5 mm long. Spores 12–13.5 µm long.

This species is characterized by short, marginal to ± marginal limbidia on the vaginant laminae of all or most leaves, pluripapillose leaf cells, and costae ending 2–3 cells below the leaf apex.

In Uganda known from a single find from Itwara FR (Kabarole District). Endemic to Africa. Reported from Benin, Central African Republic, Ghana, Ivory Coast, Nigeria, Senegal, Seychelles, Sierra Leone, and Tanzania.

Mostly on wood (bark, roots, twigs, rotten wood), less often on rock, infrequently on soil, to 1900 m.


.

Fissidens megalotis Schimp. ex Müll.Hal. subsp. helictocaulos (Müll.Hal.) Brugg.-Nann., J. Hatt. Bot. Lab. 81: 163, 1997 (Fig. 14)

Plants 4–6 X 1.5–2 mm. Leaves limbate, oblong-lanceolate, apex widely acute, often rounded mucronate, 1.25–1.35 X 0.35–4 mm, 3–4 times as long as wide. Limbidium marginal, in lower part of the vaginant laminae bordered by 1–3 rows of short, wide cells (intralaminal), reaching insertion of dorsal lamina and leaf apex, the two limbidia of the vaginant laminae not confluent distally. Vaginant laminae ½–2/3 the leaf length, slightly unequal. Dorsal lamina tapering towards below, reaching the insertion, often decurrent. Costa percurrent to excurrent. Leaf cells 7.5 X 6 µm, smooth, convex. Perichaetia terminal. Perichaetial leaves to 1.7 mm long. Perigonia and sporophytes not known from Uganda.

This subspecies is characterized by limbate leaves, slightly unequal vaginant laminae of which the limbidia not confluent distally and intralaminal below, and smooth, bulbous leaf cells. The other subspecies F. m. subsp. megalotis is known from the Cape Province (South Africa), Australia and New Zealand. It is not found in Uganda. It differs from subsp. helictocaulos in the smaller vaginant laminae ending on or near the costa and unipapillose to pluripapillose leaf cells.

Fissidens megalotis subspecies helictocaulos is endemic to Africa. In Uganda it is known from a single collection from Arua District. Further distribution: Cameroon, Cape Verde Islands, Democratic Republic of the Congo, Eritrea, Ethiopia, Kenya, Malawi, Somalia, South Africa, Tanzania, Zambia, and Zimbabwe.

Mostly on soil, less often on rotten wood or stones, humid places, frequently near streams and on road-banks, 360–2050 m.


 

Fissidens metzgeria Müll.Hal., Linnaea 39: 358, 1875 (Fig. 7d-f)

Plants 2–20 X 1.5–3 mm. Leaves limbate, elliptic-lanceolate, apex acute, ending in large pointed cell, 1.6–2.4 X 0.4–0.5 mm, 4–5 times as long as wide. Limbidium marginal, lax, reaching the insertion, ending just below the leaf apex. Vaginant laminae to ½ the leaf length, ± equal. Dorsal lamina straight to slightly rounded at base, reaching the insertion, not decurrent. Costa absent. Leaf cells large, 55–112 X 20–37 µm, flat, smooth, limbidial cells lax, 3–9 µm wide. Perigonia, perichaetia and synoecia terminal. Perichaetial leaves 2.5 mm long. No mature sporophytes seen from Uganda.

This species is characterized by limbate leaves without costa and ill-defined limbidia of long, wide cells. Limbidia of F. enervis, the only other limbate species without costa are distinct and consist of narrower, 1 µm wide, cells. Quite often F. metzgeria has clusters of multicellular gemmae on the apical part of the leaf.

In Uganda known from Bushenyi, Kabale, Kabarole, Masindi, Mpigi and Rukungiri Districts. Also known from Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko), Gabon, Guinea, Nigeria, Rwanda, and Tanzania. Endemic to Africa.

In humid places, frequently near streams, mostly on soil of road banks, less often on rotten wood or stone, to 2050 m.


.

Fissidens ovatus Brid., Muscol. Recent. 4. Suppl.: 190, 1819 (Fig. 15)

Plants 15 X 5 mm, with large axillary nodules. Leaves elimbate, lanceolate, apex acute, long acuminate to mucronate, 3.0–4.2 X 0.6–0.9 mm, 4.5–5.5 times as long as wide. Vaginant laminae ¾ the leaf length, slightly unequal. Dorsal lamina rounded-truncate, often strongly undulate at base, reaching the insertion, decurrent. Costa long percurrent to long excurrent. Leaf cells 7.5–9 X 4.5–7.5 µm, convex, smooth; marginal cells of vaginant laminae in basal part linear, in middle part short and wide, often thin-walled and empty. Perichaetia terminal on short branches. Perigonia and sporophytes not collected in Uganda.

This species is recognized by its large axillary nodules, long, elimbate leaves with excurrent costae and convex leaf cells. Also characteristic, when present, are thin-walled marginal cells on the vaginant laminae and dorsal laminae that are undulate below.

In Uganda known from Arua, Kabarole, Kapchorwa, Mbale and Moroto Districts. Endemic, common and widespread in Africa: Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko), Eritrea, Guinea, Kenya, Lesotho, Madagascar, Malawi, Mauritius, Nigeria, Réunion, South Africa, Swaziland, Tanzania, Zambia, and Zimbabwe.

In humid places, frequently in riverbeds and waterfalls, mostly on rock, also on bark, roots, rotting wood and soil, to 2850 m.


 

Fissidens pallidinervis Mitten, J. Linn. Soc. Bot. 12: 592, 1869 (Fig. 16)

Plants 1.1–1.5 X 0.5–0.6 mm. Leaves mostly elimbate, oblong to obovate, apex broadly acute to obtuse, rounded, 0.45 X 0.15 mm, 3 times as long as wide. Limbidium restricted to lower half of vaginant laminae of upper leaves of male and female plants. Vaginant laminae 1/2 the leaf length, slightly unequal. Dorsal lamina reaching the insertion to ending above in lower leaves, not decurrent. Costa ending 6–11 cells below leaf apex. Leaf cells 7.5 X 6 µm with 3–4 papillae. Perigonia and perichaetia terminal. Sporophytes not known from Uganda.

This species is characterized by broadly acute to obtuse rounded leaf tips that lack a clear apical cell, costae that end 6–11 cells below the leaf apex, pluripapillose cells and limbidia restricted to the lower half of the vaginant laminae of upper leaves of fertile stems. It resembles expressions of F. sciophyllus with a short costa. These, however, are distinct in having a clear top cell.

The Uganda specimen of F. pallidinervis has wide, multicellular gemmae in some leaf axils.

In Uganda known from one collection from Iganaga District: 10 miles (16 km) S of Bugiri, on roots, elevation unknown. Wood s.n., 1950 (BM). Further African distribution: Central African Republic, Malawi, Seychelles, Tanzania, and Zambia. Outside Africa known from the Southeastern USA, Central and South America, and Asia.

Mostly on rock, less often on bark or termite mounds, in humid to wet places from sea level to 2050 m.


 

Fissidens palmifolius (P.Beauv.) Broth. var. semilimbatus Brugg.-Nann. Cryptog. Bryol. 25: 313, 2004 (Fig. 17)

Plants 10–15 X 2 mm. Leaves semilimbate or elimbate, lanceolate, apex acute, 2.5–3.5 X 0.4–0.5 mm, 6–7 times as long as wide. Limbidium intralaminal or marginal, extending the length of the vaginant laminae, not present on all leaves. Vaginant laminae 3/5 the leaf length, unequal. Dorsal lamina narrow at base, mostly ending above insertion. Costa short, ending 15–26 cells below leaf apex. Leaf cells 12–22.5 X 7.5–21 µm, smooth and flat. Perigonia, perichaetia and sporophytes unknown.

This variety is characterized by narrow, lanceolate, 2.5–3.5 mm-long leaves, that are semilimbate on some plants and elimbate in others. F. palmifolius var. palmifolius has elimbate vaginant laminae and leaves that are about twice long. It is not known from Uganda.

Known only from the type locality: River Mpanga, Kabarole District, firmly attached to rock, presumably under water, at 1370 m (BM).


 

Fissidens pellucidus Hornsch., Linnaea 15: 146 1841 (Fig. 18)

Plants 2–5 X 1.4 mm. Leaves elimbate to weakly limbate, ovate-lanceolate, apex acute, 0.65–1.2 X 0.2–0.3 mm, 3–4 times as long as wide. Limbidium restricted to basal part of vaginant laminae of upper leaves of fertile stems. Vaginant laminae 1/2 or less as long as the leaf, unequal, lesser laminae ending halfway between margin and costa or on the costa. Dorsal lamina wide, rounded at base, reaching the insertion. Costa ending 6–11 cells below leaf apex to percurrent. Leaf cells often guttulate, 7.5–15 X 6–13.5 µm, smooth. Perichaetia terminal. Perichaetial leaves 1.2 mm long. Perigonia and sporophytes not known from Uganda.

This variable species is characterized by smooth, guttulate leaf cells. Leaves can be elimbate or weakly limbate and the costa may end far below the leaf apex or be percurrent.

Uncommon in Uganda: Arua and Rukungiri Districts, and Damba Island (Mukono). Widely distributed in Africa: Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Rio Muni, Bioko), Republic of the Congo, Gabon, Ghana, Guinea, Ivory Coast, Liberia, Madagascar, Nigeria, Réunion, Seychelles, Sudan, and Tanzania. Also known from the southern USA, Central and South America, Asia, and Australia.

Mostly on soil, also on rock and wood, termite mounds in forests. In Africa found to 2260 m.




Fissidens porrectus Mitt., Trans. Linn. Soc. London 23: 56, 1860 (Fig. 19)

Plants 6.5 x 3 mm. Leaves semilimbate, elliptic-ovate, apex acute to broadly acute, 1.8–1.9 x 0.5–0.55 mm, ± 3.5 as long as wide, margin dark, swollen, in part bistratose. Limbidium on vaginant laminae of all or most leaves, weakly intralaminal near insertion. Vaginant laminae about half the leaf length, gaping, slightly unequal. Dorsal lamina rounded below, reaching insertion, not decurrent. Costa excurrent. Leaf cells clear, often guttulate, 12–16.5 x 9–13.5 µm, smooth. Perigonia and perichaetia terminal on plants or branches. Perichaetial leaves to 2.3 mm long. Seta 5 mm long, 1–2 per perichaetium. Capsule erect, 1.1 x 0.4 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 45 µm wide at base. Operculum 1 mm long. Spores 12–15 µm.

Only one form of this variable species is known from Uganda, It is easily recognized by the dark, swollen margin of the apical and dorsal laminae, the limbate vaginant laminae and large, firm-walled, often guttulate leaf cells.

In Uganda known from Kabale, Masaka, Masindi, and Rukungiri Districts. Further distribution: Cameroon, Democratic Republic of Congo, Ethiopia, Equatorial Guinea (Rio Muni), Gabon, Guinea, Ivory Coast, Malawi, Nigeria, Rwanda, South Africa, Tanzania, and Zimbabwe. Endemic to Africa.

On rock, inundated rock, soil, living and dead wood in humid places, to 2130 m.


 

Fissidens pseudoeenii Bizot & Dury ex Pócs, Fol. Hist. Nat. Mus. Matr. 4: 30, 1977 (Fig. 20a-e)

Plants 1 x 0.7 mm. Leaves elimbate, elliptical to oblanceolate, apex acute, ending in large pointed cell, 0.45–0.55 x 0.13–0.15 mm, 3–4 times as long as wide, margin serrate, coarser serrate on vaginant laminae, entire at base of vaginant laminae. Vaginant laminae 3/5 the leaf length, unequal, lesser laminae ending halfway between margin and costa. Dorsal lamina tapering below, reaching the insertion or ending above. Costa ending 1–2 cells below leaf apex to percurrent. Leaf cells 9–10.5 x 9 µm, smooth; basal marginal 1–5 cells of vaginant laminae linear-oblong. Perigonia, perichaetia and sporophytes unknown from Uganda.

This species is characterized by small plants, elimbate leaves on infertile plants, leaf margins serrate, particularly on the vaginant laminae and large smooth leaf cells.

Uganda, Arua District. Also known from Malawi and Tanzania. African endemic.

On soil and wood, on river banks and side of an anthill, 1200–2180 m.


 

Fissidens ramulosus Mitt., Trans. Linn. Soc. London 23: 54, 1860 (Fig. 20f-k)

Plants 7–5 x 1.4–1.7 mm. Leaves mostly elimbate, oblong-lanceolate, apex acute, acuminate, ending in 1–5 large, clear cells, 0.8–1.2 x 0.2–0.4 mm, 3–3.5 times as long as wide. Limbidium marginal, restricted to the basal half of vaginant laminae of upper leaves of female plants to limbate on most leaves of fertile and some of infertile plants. Vaginant laminae ½–¾ the leaf length, slightly unequal. Dorsal lamina rounded to gradually narrowed below, reaching the insertion, not decurrent. Costa percurrent to excurrent or ending 2–3 cells below leaf apex. Leaf cells 3–7.5 x 3–6 µm, unipapillose. Perigonia axillary or terminal on plants or branches. Perichaetia terminal on plants or branches. Perichaetial leaves 1.45 mm long. Seta 2–3.5 mm long. Capsule erect to slightly inclined, 0.5 x 0.25–0.35 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 31.5–33 µm wide at base. Operculum 0.4–0.45 mm long. Spores 9–13.5 µm.

This species is characterized by elimbate to partially limbate leaves, unipapillose leaf cells and scariosus-type peristomes. It resembles F. sciophyllus which, however, has pluripapillose leaf cells. Fissidens reflexus is unipapillose and similar too, but typically has wider vaginant laminae that often have reflexed margins and a different peristome.

In Uganda rather common: Arua, Kabarole, Luwero, Masindi, Mpigi, and Rukungiri Districts. Further distribution: Angola, Cameroon, Central African Republic, Comores, Democratic Republic of the Congo, Gabon, Ivory Coast, Kenya, Madagascar, Nigeria, Rodriguez, and Tanzania. Restricted to Africa.

On rock, soil, less often on wood, bark or termite mounds, 5–1790 m.


 

Fissidens reflexus Hampe, Linnaea 38: 221, 1874 (Fig. 21a-f)

Plants often dimorphic, infertile plants 5–8 x 1.5 mm; female plants shorter, 2–4 mm long, and closer foliated. Leaves partially limbate, ovate-lanceolate to oblong, apex acute often slightly acuminate, 0.75–1.1 x 0.35 mm, 2.5–3 times as long as wide. Limbidium marginal or bordered by large, oblong cells, restricted to 1/3–3/4 of the vaginant laminae of all or most leaves. Vaginant laminae ¾ the leaf length, margins often revolute, at base wider than stem, slightly unequal. Dorsal lamina tapering to rounded-truncate at base, reaching insertion, not decurrent. Costa ending 2–3 cells below leaf apex to percurrent or excurrent. Leaf cells 6–7.5 x 4.5–6 µm, mammillose to unipapillose. Perigonia terminal, also axillary and budlike on female stems; perichaetia terminal, a few solitary, naked archegonia in leaf axils. Perichaetial leaves 1–1.2 mm long. Seta 2–2.5 mm long; capsule ± erect, 0.4–0.8 x 0.25–4 mm, ± 32 files of exothecial cells. Peristome teeth straight, undivided to irregularly divided, 33–39 µm wide at base. Operculum 0.3 mm long. Spores 16.5–22.5 µm long.

Fissidens reflexus is characterized by mammillose to unipapillose leaf cells, vaginant laminae that are mostly wider than the stem, limbidia on the lower ½–3/4 of all or most leaves of both fertile and infertile stems, and straight, short, undivided to irregularly divided peristome teeth

In Uganda known from a few finds from Arua District. Further distribution: Central African Republic, Chad, Democratic Republic of the Congo, Ivory Coast, Madagascar, Malawi, Nigeria, Seychelles, Tanzania, and Zambia. Endemic to Africa.

On earth, rock and termite mounds, 200–1650 m.


 

Fissidens robynsianus Demaret & P. de la Varde, Bull. Jard. Bot. État, 27: 757, 1957 (Fig. 21g-j)

Plants 10 x 1 mm. Leaves limbate, elliptic-lanceolate, apex acute, 1.0–1.2 x 0.25–0.3 mm, 3.5–4 times as long as wide. Limbidium marginal, ending far below the leaf apex, reaching the insertion of the dorsal lamina. Vaginant laminae 3/5 the leaf length, slightly unequal. Basal 1/5 of dorsal lamina consisting of linear cells that are not or hardly distinct from costal or limbidial cells. Costa ending 2–3 cells below leaf apex. Leaf cells 7.5–13.5 x 6–7.5 µm, smooth, ± flat. Fertile parts and sporophytes unknown from Uganda.

This species forms dense mats or tufts that are blackish with green shoots. It is recognized by its stiff, lanceolate, limbate, often blackish and worn off leaves and the basal 1/5 part of the dorsal laminae consisting of linear, pluristratose cells.

In Uganda known from Mount Elgon, Kapchorwa District. A rare African, high altitude species that is further known only from its type locality in Democratic Republic of the Congo.

Firmly attached on rocks in the spray of waterfalls and on wet rocks, 3170–3870 m.


 

Fissidens schmidii Müll.Hal., Bot. Zeit. (Berlin) 11: 18, 1853 (Fig. 22)

Plants 5–22 x 1.3–2.2 mm. Leaves limbate, lanceolate to elliptic-lanceolate, apex acute often weakly acuminate, 0.95–2.2 x 0.2–0.5 mm, 3–4.5 times as long as wide. Limbidium marginal, infrequently weakly intralaminal below, reaching the leaf apex to ending below, reaching the insertion of the dorsal lamina or not, limbidia of the two vaginant laminae mostly distally confluent. Vaginant laminae 2/3–3/5 the leaf length, equal to slightly unequal. Dorsal lamina tapering towards base, ending at insertion to long decurrent. Costa ending 1–2 cells below a leaf apex, percurrent or excurrent. Leaf cells 6–7.5 x 3–6 µm, smooth, bulging. Perigonia terminal on plants or branches; perichaetia terminal; perichaetial leaves 2.5 mm long. Seta 5 mm. Capsule erect, 0.8 x 0.4 mm, ca 50 files of exothecial cells, peristome bryoides-type, teeth 37.5 µm wide at base. Operculum 0.6 mm. Spores 12–13.5 µm.

This variable species is best recognized by its limbate, often decurrent leaves, with the limbidia of the vaginant laminae confluent above and small, bulging leaf cells. It can be confused with F. androgynus which, however, has larger, 7.5–13.5 x 4.5–10.5 µm, flat to slightly convex leaf cells. Small forms of F. leucocinctus can also be hard to separate from F. schmidii; however, leaf cells of F. leucocinctus are ± flat and not to hardly bulging.

In Uganda known from: Arua, Masindi, Kapchorwa (Mt Elgon) and Rukungiri Districts. Widespread in Africa: Cameroon, Central African Republic, Ethiopia, Gabon, Guinea, Malawi, and Nigeria. Also in Asia.

On rock, soil, bark or wood, humid to wet places, to 3870 m.


 

Fissidens sciophyllus Mitt., Trans. Linn. Soc. London 23: 55, 1860 (Fig. 23)

Plants 4–8.5 x 1.7–2 mm. Leaves elimbate or with short limbidium, lanceolate, oblong-lanceolate, apex acute, acuminate, ending in 1–5 clear cells, 1.25–1.3 x 0.27–0.4 mm, 3–4.5 times as long as wide. Limbidium marginal to intralaminal, extending ½ the length of the vaginant laminae or less of the 3 (-8) upper leaf pairs of female plants, rarely perichaetial leaves elimbate. Vaginant laminae 3/5 the leaf length, slightly unequal. Dorsal lamina slightly rounded to rounded truncate below, ending at the insertion. Costa excurrent, percurrent or ending 2–4 (–8) cells below leaf apex. Leaf cells 3–9 x 3–6 µm with 2–5 papillae. Perigonia terminal on short plants at base of female plants, on axillary branches, or budlike in leaf axils; perichaetia terminal on plants or branches. Perichaetial leaves 1.5–1.7 mm long. Seta 2–3 mm long; capsule erect, 0.6–0.7 x 0.3–0.4 mm, ± 32 files exothecial cells. Peristome scariosus-type, teeth 32.5–38.5 µm wide at base; operculum 0.4 mm. Spores 13.5–15 µm long.

This variable species is characterized by limbidia restricted to the basal half or less of upper leaf pairs of female stems, rarely all leaves elimbate, and by pluripapillose leaf cells. Leaves with a short costa end in 1 clear top cell; leaves with a percurrent costa end as 1–5 large, clear cells.

Fabronia sciophyllus fo. ulna (Müll.Hal) Brugg.-Nann. is also known from Uganda. It differs from fo. sciophyllus in limbidia that are not restricted to the perichaetial and subtending leaves, but may be present on the basal part of the vaginant laminae of most leaves. The two forms are not sharply separated, limbate expressions may have elimbate innovations. In Uganda fo. sciophyllus is the more common one.

A common species. In Uganda known from Arua, Bushenyi, Iganga, Kabarole, Kampala, Kasese, Luwero, Masindi, Mbale, Mubende and Rukungiri Districts. Further distribution: Angola, Benin, Cameroon, Cape Verde islands, Central African Republic, Chad, Democratic Republic of the Congo, Djibouti, Equatorial Guinea (Rio Muni and Bioko), Gabon, Ghana, Guinea, Kenya, Madagascar, Lesotho, Namibia, Nigeria, Réunion, Rodriguez, Rwanda, São Tomé, Seychelles, South Africa, Sudan, Tanzania, and Zambia. Restricted to Africa.

On rock, bark, roots, rotting wood, soil, and termite mounds, to 2620 m, most common below 2200 m.



Fissidens submarginatus Bruch in Krauss, Flora 29: 133, 1846 (Fig. 24a-e)

Plants 4–8 x 2 mm. Leaves semilimbate, lanceolate, apex acute, often ending in a large pointed cell, 1.5–1.85 x 0.35–0.4 mm, 4–5 times as long as wide. Limbidium marginal; near insertion infrequently intralaminal, extending the whole length of the vaginant laminae of all leaves and sometimes a short distance onto the apical lamina. Vaginant laminae 3/5–3/4 the leaf length, margins sometimes reflexed, slightly unequal. Dorsal lamina rounded or tapering below, reaching the insertion, not decurrent. Costa ending 2–4 cells below leaf apex, percurrent or excurrent. Leaf cells 4.5–7.5 x 3–6 µm, with 1 (–2) papillae. Perigonia terminal on short or long plants; perichaetia terminal. Perichaetial leaves 1.0–1.8 mm long. Seta 3.5–5 mm long, capsule slightly inclined, 0.9–1.1 x 0.4–0.5 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 34.5–46.5 µm wide at base. Operculum 0.8 mm long. Spores 9–12 µm long.

This species is characterized by completely limbate vaginant laminae and unipapillose leaf cells. For differences with F. intramarginatus see under that species. In one collection subterranean gemmae were seen.

In Uganda found in Arua, Bushenyi, Kabale, Kabarole, Kigezi, Masaka, Masindi, Mpigi, Mukono and Rukungiri Districts. Further African distribution: Cameroon, Central African Republic, Democratic Republic of the Congo, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Madagascar, Nigeria, Senegal, South Africa, Swaziland, Tanzania, Togo, Zambia, and Zimbabwe. Also known from the neotropics.

Mostly on soil, also on decaying wood, stones and termite mounds, to 2500 m.


 

Fissidens unipapillosus Brugg.-Nann., Cryptog. Bryol. 25: 309, 2004 (Fig. 25)

Plants 4–7 x 2.2 mm. Axillary nodules differentiated. Leaves limbate, undulate, oblong, apex widely acute to rounded obtuse, often apiculate, 1.3–1.8 x 0.3–0.35 mm, 4.5–5 times as long as wide. Limbidium marginal, on all laminae of all leaves, ending below leaf apex and above the insertion of the dorsal lamina. Vaginant laminae 3/4 the leaf length, margins reflexed, slightly unequal. Dorsal lamina rounded below, reaching the insertion, not decurrent. Costa excurrent to percurrent. Leaf cells 3–4.5 x 3–4.5 µm, bulging, sharply unipapillose, infrequently papillae with 2 tips. Antheridia axillary on female or separate plants, naked, 1–3 per axil, in upper axils often together with a single archegonium; perichaetia terminal. Perichaetial leaves 1.4–1.7 mm long. Seta 3–3.5 mm long, 1–2 per perichaetium; capsule inclined, 0.7 x 0.4 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 45 µm wide at base. Operculum 0.5–0.7 mm long. Spores 7.5–13.5 µm long.

This limbate species may be confused with F. weirii Mitt., which, however, has pluripapillose leaf cells. cells. Fissidens angustifolius Sull. is also unipapillose and limbate, but has larger leaf cells (7.5–10.5 x 4.5–7.5 µm) and often frondiform plants. Plants of F. unipapillosus are pinnate.

This African species is known from Uganda only: Arua, Kampala, Masindi and Mukono (Damba Island) Districts.

On damp soil, ants nest, bank of saw pit, earth in shade, on soil on top of log, 1220 m (elevation in most cases not indicated).


 

Fissidens usambaricus Broth., Bot. Jahrb. Syst. 20: 182, 1895 (Fig. 7g-i)

Plants 1.5–5 x 1.5–2.5 mm. Leaves elimbate, elliptic-oblanceolate to obovate, apex widely acute to obtuse and rounded, often weakly apiculate, 1.0–1.5 x 0.33–0.6 mm, 2.5–3 as long as wide. Vaginant laminae to half as long as the leaf, slightly unequal. Dorsal lamina slightly rounded below, reaching the insertion, sometimes slightly decurrent. Costa absent except for a weak costa extending a few cells above its departure from the vaginant laminae in some perichaetial leaves. Leaf cells 42–75 x 15–36 µm, smooth, flat to slightly convex, marginal cells smaller, 12–30 x 7.5–10.5 µm. Perigonia, perichaetia and synoecia terminal. Perichaetial leaves 2.2 mm long. Seta 2.5–3 mm long, 1–3 per perichaetium; capsule erect to slightly inclined, 0.65 x 0.45 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 37.5–45 µm wide at base. Operculum 0.5–0.55 mm long. Spores 9–12 µm long.

This species is characterized by elimbate, ± costa-less leaves and very large leaf cells. Fissidens metzgeria and F. enervis which also lack costae and have with large cells, have limbate leaves.

In Uganda collected in Kabale, Kabarole, Masindi, Mukono (Damba Island) and Rukungiri Districts. Further African distribution: Cameroon, Cape Verde Islands, Central African Republic, Democratic Republic of the Congo, Kenya, Nigeria, Rwanda, South Africa, Swaziland, and Tanzania. Endemic to Africa.

Mostly on soil, rarely on wood or rock, 900–2400 m.


 

Fissidens weirii Mitt., J. Linn. Soc. Bot. 12: 602. 1869 (Fig. 24f-k)

Plants 3.5–10 x 2.2–2.5 mm wide. Leaves limbate, lanceolate to oblong lanceolate, undulate, apex acute to rounded-obtuse with apiculus, 1.5 x 0.3 mm, ± 5 times as long as wide. Limbidium marginal, on all laminae of all leaves, ending above insertion of the dorsal lamina and below leaf apex. Vaginant laminae ¾ the leaf length, margins often reflexed, slightly unequal. Dorsal lamina rounded at base, reaching the insertion, not decurrent. Costa percurrent to excurrent. Leaf cells 4.5–6 x 3–4.5 µm, with 1–4 papillae. Perigonia axillary, budlike, infrequently the buds are synoecia; perichaetia terminal. Perichaetial leaves 1.5 mm long. Seta 6 mm long; capsule slightly inclined to erect, 0.9–1 x 0.4–0.5 mm, exothecial cells in ± 32 files. Peristome scariosus-type, tooth base 45 µm wide. Spores 9–13.5 µm long.

This species is characterized by pluripapillose leaf cells, limbate leaves and limbidia ending below the leaf apex and above the base of the dorsal lamina. Infrequently leaves vary from limbate on all laminae to limbate on the vaginant laminae only in one and the same collection. Fissidens weirii differs from the also limbate, pluripapillose F. glaucissimus in its wider leaves and narrower limbidia on the vaginant laminae.

In Uganda known from Masaka and Rukungiri Districts. Further African distribution: Angola, Burkina Faso, Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Rio Muni, Bioko), Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Malawi, Nigeria, Sierra Leone, and Togo. Also widespread in the neotropics.

In Africa found on termite mounds, rock, soil and wood, sea-level to 2460 m.


 

Fissidens zollingeri Mont., Ann. Sci. Nat. Bot. 4:86: 114, 1845 (Fig. 26)

Plants 1.3–5 x 1–3.5 mm, often flabelliform. Axillary nodules often differentiated. Leaves limbate, elliptico-lanceolate, apex acute to acute-acuminate, upper leaves 1.2–2.5 x 0.3–0.5 mm, 4–7 times as long as wide. Limbidium marginal, on all laminae of all leaves reaching the leaf apex and nearly to completely reaching the insertion. Vaginant laminae 3/5–2/3 the leaf length, equal. Dorsal lamina narrowed below, reaching the insertion in upper leaves to ending above in lower ones, not decurrent. Costa percurrent to excurrent. Leaf cells 7.5–15 x 4.5–9 µm, flat to slightly convex; in vaginant laminae often patches of clear, 18–42 x 12–15 µm, cells. Perigonia terminal on plants at base of female stems; perichaetia terminal. Perichaetial leaves 1.5–3 mm long. Seta 3.5–6 mm long; capsule erect, 0.7 X 0.3 mm, ± 32 files of exothecial cells. Peristome scariosus-type, teeth 37.5 µm wide at base. Operculum 0.8 mm long. Spores 9–12 µm long.

This species can be confused with F. curvatus, which, however, has more than 40 files of exothecial cells, a bryoides type peristome and dimorphic stems. Fissidens angustifolius Sull. too is similar, but has unipapillose laminal cells.

In Uganda known from: Arua, Kabarole, Kampala, Luwero, Masaka, Masindi and Mpigi Districts. Further African distribution: Angola, Benin, Cameroon, Comores, Democratic Republic of the Congo, Equatorial Guinea (Bioko), Gabon, The Gambia, Guinea, Kenya, Madagascar, Nigeria, Rodriguez, Seychelles, and Tanzania. Outside Africa it is known from the neotropics, Asia and Oceania.

On soil, wood and rarely termite mounds, 0–3350 m.


 

Excluded taxa

Fissidens borgenii Hampe
The two specimens cited by Porley et al., (1999) as F. borgenii Hampe are F. sciophyllus Mitt.

Fissidens erosulus (Müll.Hal.) Paris
All Ugandan specimens identified as F. erosulus seen by me appeared to be either F. ramulosus or F. sciophyllus Mitt.

Fissidens crispulus Brid.
The occurrence of F. crispulus in Uganda could not be confirmed. Tracing back the citation by O’Shea (1999) it appeared that F. crispulus has never been reported from Uganda. O’Shea (1999) was based on Kis’ citation (1985) of F. sylvaticus Griff. Most African specimens of F. sylvaticus from Africa are indeed either F. ovatus Brid. or F. crispulus Brid. However, those cited by Kis are not. She based herself on Potier de la Varde (1955, 1956) who reported F. amblyophyllus Müll.Hal. from Uganda. Kis’ translation of F. amblyophyllus into F. sylvaticus is easily explained: Bizot & Pócs (1979) reduced F. amblyophyllus to F. sylvaticus Griff. and Kis (1985) must have overlooked that Magill (1981) corrected this and reduced F. amblyophyllus Müll.Hal. to F. asplenioides Hedw. I checked several specimens identified as F. amblyophyllus by Potier de la Varde including Wood 1243 from Uganda (BM) cited by Potier de la Varde (1956). All proved to be F. asplenioides Hedw.

Fissidens palmifolius (P.Beauv.) Broth. var. palmifolius.
Pursell (1987), based on a single specimen (Thomas s.n. (Dixon 1423-b) (BM)), cited this species from Uganda. This specimen, however, appeared to differ from F. palmifolius var. palmifolius in having leaves half as long, and that are also often semilimbate instead of elimbate. These differences were considered large enough to warrant describing a new variety: F. palmifolius var. semilimbatus.


 

References:

Bizot M, Pócs T. 1979. East African Bryophytes, III. Acta Botanica Academiae Scientiarum Hungaricae 25 (3,4): 223-261.

Bruggeman-Nannenga MA, Berendsen W. 1990. On the peristome types found in the Fissidentaceae and their importance for the classification. Journal of the Hattori Botanical Laboratory 68: 193–234.

Iwatsuki Z, Suzuki T. 1982. A taxonomic revision of the Japanese species of Fissidens (Musci). Journal of the Hattori Botanical Laboratory 51: 329-508.

Kis G. 1985. Checklist of the mosses of South-east Tropical Africa. Proceedings of the third meeting of the Bryologists from Central and East Europe, Praha, 14th-18th June 1982, Univerzita Karlova Praha 1984: 217-282.

Magill RE. 1981. Flora of Southern Africa. Bryophyta Part I. Mosses. Fasc. I. Sphagnaceae–Grimmiaceae. In: Leistner OA, Flora of Southern Africa. South Africa: Botanical Research Institute, Department of Agriculture and Fisheries.

O'Shea BJ. 1999. Checklist of the mosses of sub-Saharan Africa, version 3. Tropical Bryology Research Reports 1: 1-133.

Porley RD, O'Shea BJ, Wigginton MJ, Matcham HW, Hodgetts NG, Stevenson CR. 1999. Bryophytes of Uganda 2. New and interesting records. Tropical Bryology 16: 179–194.

Potier de la Varde R. 1955. Mousses récoltées par M. le Dr. Olov Hedberg, en Afrique orientale, au cours de la Mission suédoise de 1948. Arkiv för Botanik 3: 125-204.

Potier de la Varde R. 1956. Contribution à la flore bryologique africaine (huitième article). Revue Bryologique et Lichénologique 25: 213-233.

Pursell RA. 1987. A taxonomic revision of Fissidens subgenus Octodiceras (Fissidentaceae). Memoirs of the New York Botanical Garden 45: 639-660.