|
| 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. |
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 unmistakable
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.
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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.
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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.
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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).
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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.
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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.
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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.
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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.
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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.
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