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Bothalia 27,1: 17-27(1997)
Studies in the liverwort genus Fossombronia (Metzgeriales) from southern
Africa, 1. Three new species from Northern Province, Gauteng and Mpu
malanga
S.M. PEROLD*
Keywords: Fossombronia, Fossombroniaceae, F. gemmifera sp. nov., F. glenii sp. nov., F. straussiana sp. nov., liverworts, Metzgeriales
ABSTRACT
Three new species of Fossombronia from Northern Province, Gauteng and Mpumalanga (formerly Transvaal) are described:
F. gemmifera, F. glenii and F. straussiana. F. gemmifera is distinguished by a highly convoluted pseudoperianth, the frequent
presence of gemmae produced dorsally on the stem, lamellate spores and mostly rather short elaters; F. glenii can be recognized
by a smallish, deeply lobed pseudoperianth, perigonial bracts with finger-like projections shielding the antheridia and by spinous
spores; F. straussiana is distinct by its hyaline or brownish rhizoids, by its dense, frilly leaves, its pseudoperianth with lamellate
lateral outgrowths, by spore ornamentation that usually has inclusions in the incomplete areolae and by the elaters which are
finely papillose.
INTRODUCTION
Within the order Metzgeriales the cosmopolitan sim-
ple-thalloid liverwort genus Fossombronia Raddi is clas
sified in the subfamily Fossombronioideae Engl, emend.
R.M.Schust. (family Fossombroniaceae Hazsl. (= Co-
doniaceae H.Klinggr.). The Fossombroniaceae is the only
family under the suborder Fossombroniineae R.M.Schust.
and is regarded as phylogenetically pivotal (Solomon
1995). It comprises four genera: Petalophyllum Nees &
Gottsche ex Lehm., Sewardiella Kashyap, Austrofossom-
bronia R.M.Schust. and Fossombronia, of which only the
latter is known to occur in southern Africa.
Plants of the genus Fossombronia are small, usually
gregarious with mostly prostrate, fleshy stems, laterally
bearing succubously inserted, undulating or crisped thallus
wings, dissected into leaf-like segments and free to the
base, where they are bistratose, but unistratose elsewhere.
The stems are anchored to the substrate by rhizoids which
are usually purple, but hyaline in a few rare species.
The gametangia develop acropetally and are situated
dorsally along the stem between the leaf insertions. They
are either intermingled or borne on separate shoots in
monoicous species, or they may occur on separate plants
in dioicous species. The antheridia are short-stalked, ±
globose and either naked or shielded by perigonial bracts.
The archegonia are exposed, but after fertilization the de
veloping sporophyte becomes surrounded by a campanu-
late pseudoperianth, constricted at the sometimes stipitate
base, and flaring at the plane, lobulate or crispate mouth.
The globose capsule is raised on a moderately short or
long seta, 6 or 7 to 10 cells in diameter; the capsule wall
is bistratose, the outer wall with delicate, hyaline cells and
the inner wall with irregularly quadrangular cells, contain
ing nodular and semi-annular thickenings. The walls rup
ture irregularly into small plates, releasing the spores and
* National Botanical Institute, Private Bag X101, Pretoria 0001.
MS. received: 1996-06-25.
elaters. The spores are relatively large and highly ornamented
with spines, lamellae or areolae. The ornamentation on the
outer face is generally considered species-specific and re
garded as essential for identification, although ‘so variable
that patterns of ornamentation . .. must be used to define
taxa’ (Schuster 1992). The elaters may be well or occasion
ally poorly developed and are usually 2- or 3-spiral. The
determination of sterile material is extremely difficult, if not
impossible because of the plasticity of the gametophytes, the
morphology of which varies considerably according to en
vironmental conditions.
HISTORICAL NOTES
The genus Fossombronia has been relatively poorly
studied in southern Africa. Lehmann (1829) described
Fossombronia (sub Jungermannia) leucoxantha, collected
by Ecklon on Table Mountain, Devil’s Peak and Lion’s
Head and also reported the presence of F. (sub Junger
mannia) pusilla L. on Table Mountain. Gottsche et al.
(1844-1847) confirmed the presence of F. pusilla Nees
and described F. crispa Nees from Promontorio Bonae
Spei. Krauss (1846) reported F. angulosa ‘in rupibus ri-
vulor montium natalensium’. Mitten (1878) described F.
tumida collected by the Rev. A.E. Eaton at the foot of
Lion’s Head and endorsed the records of F. crispa and F.
leucoxantha from localities near Cape Town. Stephani
(1900) described F. zeyheri and F. spinifolia. Sim (1926)
added no new species, but Amell (1952) described three
from the Cape: F. capensis, F. densilamellata and F. mon-
taguensis (Perold 1997: 29). Amell (1963) stated that ‘the
South African species of the genus are in great need of
revision’. Scott & Pike (1984, 1987a-c, 1988a, b), who
studied the genus in Australia (i.e. the western parts of
the State of Victoria and adjacent regions of South Aus
tralia and New South Wales) for six years and described
many new species from there, expected a comparable spe
cies richness to emerge from a long-term study of the
genus in South Africa (and South America), since they
regarded its origins to have been mainly in Gond-
wanaland.
18 Bothalia 27,1 (1997)
FIGURE 1.— Fossombronia gemmifera. A-D, leaves; E, opened pseudoperianth; F, pseudoperianth from side; G-J, perigonial bracts; K, cross section
of stem; L, detail of marginal area of leaf, with slime papilla (see arrow); M, median leaf cells with oil bodies and chloroplasts; N, cells in
capsule wall. A, D, Strauss & Relief CH13655; B, C, Perold & Koekemoer 31 16b\ E, F, Perold & Van Rooy 3559a\ G-N, Strauss 132. Drawn
by G. Condy. Scale bars: A-F, 500 pm; G-J, 250 pm; K, 100 pm; L-N, 50 pm.
Bothalia 27,1 (1997) 19
This paper is the first of a series dealing with southern
African Fossombronia species.
METHODS
Samples of field-collected specimens were transferred
to a conically shaped fine-mesh sieve and washed in a jet
of running water to clear away soil particles. Remaining
particles were manually removed by using fine-tipped for
ceps. Cross sections were cut of some cleaned stems; several
leaves and a pseudoperianth were detached and all trans
ferred to a drop of water on a clean slide. Finally a coverslip
was applied to the preparation, which was stored in a covered
plastic dish lined with damp filter paper. During examination
of the slide the evaporated water was periodically replen
ished. The structures were measured and also photographed
under a compound light microscope.
The remaining portion of the cleaned specimen was
fixed in FAA (formaldehyde/alcohol/glacial acetic acid
and distilled water in proportion of 2:1:1:20). For later
reference some thalli were permanently preserved in FAA,
the remainder only for several hours and then dehydrated
in an ascending series of acetone to 100% and critical
point dried in a Balzers Union dryer, using liquid CO2 as
the transitional fluid. The thalli were mounted on alu
minium stubs with double-sided Sellotape, gold-coated,
then viewed and photographed, using an ISI SX 25 scan
ning electron microscope (SEM).
There are some advantages to studying and photo
graphing Fossombronia material, treated as described
above, with the aid of SEM. Scott & Pike (1984) stated
that the form, colour and orientation of the leaves (of dif
ferent species) are distinct but ‘beyond our powers to il
lustrate and exceedingly difficult to describe’. Although
the colour cannot be recorded by SEM, the form and cer
tainly the orientation of the leaves can. Scott & Pike
(1984) also observed that “male plants have often shriv
elled and disappeared by the time the corresponding
spores are ripe and the gametophyte is then so desiccated
that its pristine vegetative appearance is irrecoverable’.
Samples taken at different stages and treated as above,
would have made comparisons easier for them. Scott &
Pike did not illustrate the plants of any of their new spe
cies, except F. rudis (Scott & Pike 1988b) and relied heav
ily on descriptions and the spore ornam entation to
distinguish between species.
The spores and elaters were mounted on slides in
Hoyer’s fluid for examination and measurement by LM.
For SEM microscopy some spores and elaters from the
same capsule were allowed to air-dry, mounted on stubs
with double-sided Sellotape, gold coated, viewed and
photographed. SEM m icrographs of Fossombronia
spores have frequently been published and have been
used as an aid to identification. Features of spore mor
phology should be used with caution, however, because
they can be rather variable, and the spores need to be
fully mature. The Degree Reference System was used
for recording distribution data (Edw ards & Leistner
1971).
1. Fossom bronia gemmifera Perold, sp. nov.
Plantae repentes, gregariae vel dispersae; saepe gem-
mas dorsaliter secus caulem ferens. Folia imbricata, fim-
briata, apicem versus aliquando longiora quam latiora,
basin versus plerumque breviora quam latiora, nonnulla
leviter bilobata, alia apicem versus undulata. Rhizoidea
purp urea. M onoicae, interdum ut v idetur dioicae.
Antheridia bracteis tecta. Pseudoperianthium breviter
setatum, orificio intricate convoluto, in lobis multis di-
viso. Sporae 55.0-62.5 (im diametro, superficie distali
lamellis minimum 14 irregularibus duplicato-parietatis,
aliquando areolas imperfectas facientibus; superficie
proximali sine nota triradiata distincta, areolis imperfec-
tis, parietibus altis circumcinctis. Elateres 100-160 pm
longi, medio 7.5 |im lati, ex tremitates versus decre-
scentes, bis vel ter spirales, sed interdum arete spirales,
50.0-62.5 |jm longi, 12.5 |jm lati.
TY PE .— Northern Province, 2427 (T hab azim bi):
Kransberg, Farm Geelhoutbosch, on streambank, directly
south of rondavel, (-BC), S. Strauss 132 (PRE, holo.).
2529 (Witbank): ± 80 km E of Pretoria, on PretoriaAVit-
bank road, facing Balmoral turnoff, left side of road at
seepage area, (-CC), Perold & Van Rooy 3559a (PRE.
para.).
Plants smallish to medium-sized, creeping, gregarious
or scattered, green, proximal leaves sometimes clasped
around stem, occasionally with pink margins; shoots sim
ple, 5.0-9.0 mm long, 0.85 mm high, 2.6 mm wide, or
once/twice furcate, apical segments moderately divergent
(Figure 2A) 2.0-5.0 mm long, basally ± 3 mm long. Stems
prostrate, in cross section apically 300-325 pm (12 or 13
cell rows) high, 400-500 pm wide, basally 250 x 300 pm,
plano-convex (Figure IK). Rhizoids purple, 12.5-20.0 pm
wide, some with internal mycorrhizal hyphae. Leaves
overlapping, frilly, obliquely inserted succubously, vari
ously shaped, toward apex sometimes longer than wide,
but more proximally usually shorter than wide, 925-1525
x 1325-1625 pm, some slightly bilobed (Figure 1A, D),
others ruched above (Figure IB, C); lateral margins with
3 or 4 uni- or bi-celled slime papillae (Figure 1L). Leaf
cells thin-walled, at upper margins subquadrate to rectan
gular across, 32.5-50.0 x 22.5-30.0 pm, at lateral margins
long-rectangular, up to 75 x 30 pm; upper laminal cells
(4-)5- or 6-sided or polygonal, 32.5-50.0 x 22.5-30.0 pm,
middle laminal cells 62.5-72.5 x 37.5-50.0 pm, basal
cells 112.5-137.5 x 37.5-62.5 pm. Oil bodies glistening,
13-36 per cell, rounded, up to ± 3.5 pm in diameter; chlo-
roplasts numerous, round or oblong, 5.0-7.5 pm in diame
ter (Figure 1M).
?Monoicous, some specimens with shoots bearing both
antheridia and archegonia, b ut oc casionally only
antheridia or often only archegonia present. Antheridia
dorsal on stem, interspersed between archegonia, short-
stalked, globose, ± 185 pm in diameter, shielded by an
irregularly shaped perigonial bract (Figure 2C), 500-700
x 380^ 50 pm, margins toothed, with projecting cells and
slime papillae (Figure 1G-J), cells in interior 4-6-sided,
80.0-90 .0 x 3 0.0-37 .5 pm; som etimes antheridia in
groups between leaves and then lacking bracts or with
reduced bracts (Figure 2B). Archegonia numerous along
stem, naked, up to 3 per shoot becoming fertilized, soon
20 Bothalia 27,1 (1997)
FIGURE 2.— Fossombronia gemmifera. A, stem branching near apex, bracts and pseudoperianth with capsule shown; B, antheridia between leaves;
C, detail o f perigonial bracts with mostly obscured antheridia and young fertilized archegonia; D, convoluted pseudoperianth from above; E,
archegonia and gemmae between leaves; F, close-up of gemmae. A, C, D, Strauss 132; B, Perold & Koekem oer 3116b\ E, F, Perold & Van
Rooy 3557. A, x 13; B, x 26; C, x 21; D, E, x 20; F, x 54.
forming young pseudoperianths. Pseudoperianths often
crowded together, in acropetal sequence, raised on a short
stalk, ± 210 x 470 pm, widely flaring above (Figure IF),
± 900 pm long, up to 2000 fim wide across intricately
convoluted mouth (Figure 2D), divided into numerous
lobes (Figure IE), 200-500 x 150-440 pm, some apically
rounded, others with an acute apex ending in a papilla;
cells comparable in shape and size to those of leaves.
Capsules globose (Figure 2D), up to ± 600 pm in diame
ter, enveloped in a calyptra which is later shed, capsule
wall bistratose, cells in inner layer irregularly shaped,
27.5-50.0 x 25.0-37.5 pm, each cell wall with (1)2-4
nodular and occasionally semi-annular thickenings (Figure
IN). Seta delicate, 1.2-3.2 mm long, 270-350 pm in di
ameter, 6 cells across. Spores brown, hemispherical (Fig
ure 3C), 55.0-62.5 pm in diameter, including lamellae
projecting around circum ference, distal face convex
(Figure 3A, B), with at least 14 irregular, mostly double
walled lamellae running across, ± 7.5 pm apart, occasion
ally anastomosing and forming incomplete areolae;
proximal face (Figure 3D, E) lacking distinct triradiate
mark, ornamentation seemingly ‘raised’ from an encir
cling, marginal furrow (Figure 3E), tall walls forming
small, irregular, incomplete areolae, up to 5 pm wide,
around periphery 19-22 projecting ‘ends’ of lamellae, ±
5 pm long. Elaters (Figure 3F) yellow-brown, smooth,
mostly rather small and som etimes poorly formed,
(50.0-)62.5-85.0 pm long, medianly 12.5 pm wide, 3(-4)-
spiral and loosely coiled, tapering toward tips, 5 pm wide
and ending in bispiral loops, rarely 100-160 pm long, 7.5
pm wide medianly, 3-spiral. Vegetative reproduction by
gemmae (Figure 2E, F), abundantly borne dorsally along
stem, short-stalked, spindle-shaped, up to 300 x 170 pm,
green when fresh, turning brown with age.
Fossombronia gemmifera grows on sandy loam soil on
stream banks or at seepages, often mixed with mosses or
other Fossombronia species, particularly F. straussiana,
which has hyaline rhizoids and is thus easily distinguished
from it. This new species is quite distinctive with its
highly convoluted pseudoperianth and by the frequent
presence of dorsal gemmae for which it has been named.
Its spore ornamentation with double-walled lamellae on
the distal face and ‘raised’ on the proximal face above a
clear marginal furrow, is also unique. There are two other
species with purple rhizoids from the same distribution
area: F. glenii which has spinous spores and F. zeyheri
which has reticulate spores. Fossombronia gemmifera has
been collected at several localities in Northern Province
and Gauteng (Figure 4).
2. Fossombronia glenii Perold, sp. nov.
Plantae repentes, gregariae vel crebrae in coloniis. Fo
lia imbricata, undulata, valde obliquiter inserta, plerumque
oblonga, longiora quam latiora, apicem versus latiora
quam basi, apice subtruncata, lobis vadosis, angulatis. Rhi-
zoidea purpurea. Dioicae. Antheridia bracteis cum laciniis
digitiformibus tecta. Pseudoperianthium infundibulifor-
me, orificio in lobis pluribus diviso, aliquot ad basin di-
visis. Sporae 40.0-52.5 pm diametro, superficie distali
cum spinis multis altis conicis, raro fractis vel cristis bre-
vibus junctis; superficie proximali sine nota triradiata
distincta, cum tuberculis multis tenuibus vel grossis in-
spersis. Elateres 70.0-137.5 x 7.5-10.0 pm, bis vel ter
spirales.
TYPE.—Northern Province, 2427 (Thabazimbi): Water-
berg, Welgevonden Estate, cliffs at drift over Sterkstroom
above farmhouse, in partial shade, with Fissidens erosulus
(Mull.Hal.) Paris, (-BD), H.F. Glen 2146 (PRE, holo.);
H.F. Glen 2134, same locality (PRE, para.).
Bothalia 27,1 (1997) 21
FIGURE 3.—Fossombronia gemmifera. Spores and elater. A, B, distal face; C, side view o f distal face; D, E, proximal face; F, elater. A, B, D, S.M.
Perold 2017 ; C, E, F, Strauss 132. A, x 734; B, D, x 688; C, E, F, x 535.
Plants m ediu m-sized, creeping, gregarious or in
crowded colonies, green; shoots mostly simple, 5.0-7.0
mm long, 1.8 mm high, 3.0 mm wide, occasionally once
furcate close to apex (Figure 6A) or to base (Figure 6B),
moderately to widely divergent, apical segments 2-5 mm
long. Stems prostrate, fleshy, in cross section at apex
400-500 (im (13 cell rows) high, 530-650 jim wide, ta
pering proximally, at base (300-) 400-450 x 400-480 |im,
plano-convex (Figure 51). Rhizoids purple, 12-20 jim
wide. Leaves overlapping, undulating, very obliquely in
serted succubously, generally oblong, longer than wide
and wider above than basally, 2000-2500 |im long, width
1325-2500 |im above, 1050-1400 ^im below, apex sub-
truncate, with shallow angular lobes (Figure 5A-F); mar
gins with or without 1 or 2 slime papillae, the lower one
on proximal edge of leaf below midline often 2-celled
(Figure 5N). L eaf cells thin-walled, at upper margins
subquadrate, 25-30 x 32-45 jim, at lateral margins long-
rectangular, up to 87.5 x 22.5 jim; upper laminal cells 5-
or 6-sided, 57.5-75.0 x 27.5-37.5 ^m; middle laminal
cells 62.5-102.5 x 42.5 x 47.5 pm; basal cells 75.0-87.5
x 37.5-40.0 |im. Oil bodies glistening, numerous, more
than 50 per cell, up to 2.5 |im in diameter; chloroplasts
densely scattered in cells when fresh, but later tending to
clump together, ± 5 |im in diameter (Figure 50).
Dioicous. Male plants hardly smaller than females.
Antheridia dorsal between leaves, in a row, white when
immature, later turning yellow, globose (Figure 6C), short-
stalked, ± 210 |im in diameter, posteriorly shielded by
perigonial bracts (Figure 5J-M), ± 550 x 370 jim, cells
in interio r subquadrate to hexagonal, 45.0-6 2.5 x
40.0-42.5 ^m, apices divided into (2)3 or 4 finger-like
projections (Figure 6C), 150-300 |im long and 1 or 2 cell
rows wide. Archegonia naked, in an interrupted, irregular
row or in groups dorsally along the stem (Figure 6D).
Pseudoperianths produced in acropetal sequence, 1 or 2
per shoot, the younger one near the apex and the other
(if present) usually more proximally, but occasionally in
close proximity; sessile, ± funnel-shaped (Figure 5H), at con
stricted base 425-525 |im wide, up to 1750 jim long, width
across flaring mouth 2250-2375 |im, consisting of (4)5 or 6
lobes (Figure 6E), with 1^4(5) deep clefts to near the base
(Figure 5G), apices truncate, entire, or with sharply pointed
projections; cells comparable in shape and size to those of
leaves. Capsules globose, ± 625 |jm in diameter, initially
entirely enveloped in calyptra, which is later shed, revealing
capsule wall (Figure 6F), the latter bistratose, the inner cell
layer with irregularly quadrangular cells, 32.5-37.5 x
20.0-37.5 |im, at each cell wall 2 or 3 nodular and sometimes
semi-annular thickenings, (Figure 5P). Seta delicate, up to
5.6 mm long, 110 |jm in diameter. Spores brown, ± hemi
spherical (Figure 7D), 40.0-52.5 |im in diameter; distal face
convex, usually covered with numerous tall, conical spines
up to 5 (im long (Figure 7A), rarely broken (Figure 7B), and
FIGURE 4.— Map showing distribution of F. gemmifera, □; F. glenii, • ;
and F. straussiana, O, in Northern Province, Gauteng and Mpu
malanga.
22 Bothalia 27,1 (1997)
FIGURE 5.— Fossombronia glenii. A-F, leaves; G, opened pseudoperianth; H, pseudoperianth from side; I, stem in cross section; J-M, perigonial
bracts; N, detail of marginal area of leaf, with slime papilla (see arrow); O, median leaf cells with oil bodies and chloroplasts; P, cells in
capsule wall. A-C , E-K, M-P, S.M. Perold 3052\ D, L, Glen 2134. Drawings by G. Condy. Scale bars: A-H, 500 pm; J-M, 250 pm; I, 100
pm; N-P, 50 (jm.
Bothalia 27,1 (1997) 23
FIGURE 6.—Fossombronia glenii. A, stem branching near apex; B, stem branching near base; C, antheridia shielded by perigonial bracts; D,
archegonia naked along stem, young pseudoperianth near apex; E, pseudoperianth with capsule from above; F, close-up of capsule in
pseudoperianth. A, Perold & Van Rooy 3569\ B-E, Perold & Van Rooy 3568\ F, Glen 2134. A, B, D, E, x 7; C, x 15; F, x 23.
sometimes connected by short ridges, even occasionally
forming some high-walled areolae (Figure 7C); proximal
face ± flat, lacking a distinct triradiate mark, sprinkled
with numerous fine to coarse tubercles (Figure 7E), which
are sometimes rather flattened; circumference with numer
ous, up to 45, projecting spines. Elaters yellow, smooth,
70.0-137.5 \im long, 7.5-10.0 |im wide in middle and
tapering slightly toward tips, 2- (Figure 7F) and some
times 3-spiral in the same elater.
Fossombronia glenii grows on rather sandy soil in dry
stream beds or on stream banks, or else on soil pockets
in exposed rocky cliffs above streams. So far, it has only
been collected at a few localities in Northern Province,
Gauteng and Mpumalanga (Figure 4), but it is surely more
widespread. It is easily recognized by the rather small
pseudoperianth split into several lobes, by perigonial bracts
with finger-like projections and by spinous spores. Because
of its spinous spores, a specimen collected by Mogg (CH
FIGURE 7.— Fossombronia glenii. Spores and elater. A-C, distal face; D, side view of distal face; E, proximal face; F, elater. A, F, Glen 2146 ’, B,
D, Mogg C HI57; C, S.M. Perold 3052; E, Glen 2134. A, x 562, B, x 700; C, x 737; D, x 694; E, x 582; F, x 575.
24 Bothalia 27,1 (1997)
157) at Wonderboom Poort, Pretoria, was previously misi-
dentified as the so-called F. crispa, but the name had been
misapplied (Perold b in press). Fossombronia glenii is dis
tinguished from the winter rainfall species, F. leucoxantha,
by the dentate leaves and pseudoperianths (Perold a in press).
Both species have spinous spores.
This species is named in honour of an esteemed col
league at the National Botanical Institute, Dr H.F. Glen,
who has often collected liverworts, together with his wife,
Mrs R. Glen (another colleague) and their young daughter,
Melissa.
3. Fossombronia strau ssiana Perold, sp. nov.
Plantae repentes, crebrae in coloniis. Folia plerumque
valde convoluta, dense imbricata, irregulariter formata,
longiora vel breviora quam latiora, interdum cum appen-
diculo oblongo in basi proximali. Rhizoidea hyalina vel
brunnescentia. Monoicae. Antheridia archegoniaque con-
ferta, appendiculo foliari basali partialiter circumdata.
Pseudoperianthium campanulatum, orificio patenti lobati,
processibus plures lamellatis lateraliter procurrentibus.
Sporae 35.0-42.5 fim diametro distaliter usque ad 10
lam ellis dis contin uis, cristis ten uibus in terjunctis,
aliquando areolas imperfectas facientibus, saepe inclu-
sionibus papilliformibus vel cruciformibus; superficie
proximali cum nota triradiata imperfecta, non valde
distincta, superficiebus cum papillis humilibus et cristis
irregularibus brevibus tectis. Elateres 107.5-175.0 pm
longi, ter vel bis spirales, papillis tenuibus humilibus tecti.
TYPE.— Northern Province, 2427 (Thabazimbi): Krans-
berg, Farm Geelhoutbosch, on streambank, directly south
of rondavel, (-B C), S. Strauss 133 (PRE, holo.); S. Strauss
134, same locality (PRE, para.).
Plants medium-sized, creeping, in crowded colonies,
green; shoots simple, up to 9.5 mm long, 1.4 mm high,
2.3-3.0 mm wide, or once-furcate, apical segments nar
rowly divergent (Figure 9A), 4.5-7.5 mm long, basal part
3.0-4.5 mm long. Stems prostrate, fleshy, in cross section
at apex (300-)400-500 pm (13 cell rows) high, 550-680
pm wide, tapering basally to 300 x 320 pm, plano-convex
(Figure 8K). Rhizoids of all plants entirely hyaline or
brownish, 10.0-12.5 pm wide. Leaves mostly highly con
voluted, densely imbricate (Figure 9B), obliquely inserted
succubously, irregularly shaped, longer than wide (Figure
8B, C) to shorter than wide (Figure 8A, E, F), (575-)
1300-1500 x (850-) 1025-1500 pm, sometimes with an
oblong appendage at proximal base (Figure 8C, D), ± 675
x 500 pm; margins with up to 8 slime papillae at angu
lations or in between, 22.5-27.5 x 22.5-25.0 pm. Leaf
cells thin-walled, at upper margins subquadrate or rectan
gular across (Figure 8L), 25.0-27.5 x 30.0-40.0 pm; at
lower lateral margins long-rectangular, up to 47.5 x 22.5
pm; u pper lam inal cells subquadrate, 25.0 -32.5 x
30.0 -32.5 pm; middle laminal cells 5- or 6-sided,
47.5-62.5 x 27.5-40.0 pm; basal cells 70.0-75.0 x 50.0
pm. Oil bodies glistening, faintly granular, 10-25 per cell,
up to 2.5 pm in diameter; chloroplasts densely scattered
in cells, 3-5 pm in diameter (Figure 8M).
Monoicous. Antheridia short-stalked, globose, 110-135
pm in diameter, dorsal on stem between leaves, basal leaf
appendage partly curved around 1 or 2 (Figure 9D, E),
sometimes leaf appendage detached, forming an oblong
perigonial bract (Figure 8J), 430-620 x 300-420 pm, mar
gin with 1 or 2 papillae. Archegonia in close proximity
to antheridia. Pseudoperianths soon forming (Figure 9C)
after fertilization in acropetal sequence, often 2 per shoot,
close together near apex, older more proximal one some
times with capsule already dehisced; sessile, campanulate
(Figure 9F), at constricted base ± 600 pm wide, 1250 pm,
rarely to 1825 pm long, width across flaring mouth 2250
pm, margin ± scalloped, consisting of 4-7 shallow,
rounded lobes (Figure 8H), several lamellate outgrowths
projecting laterally from sides (Figure 81); cells compara
ble in shape and size to those of leaves. Capsules globose,
500-610 pm in diameter, capsule wall bistratose, cells in
inner layer irregularly shaped, 30.0-37.5 x 15.0-20.0 pm,
crowded with nodular and some semi-annular thickenings
(Figure 8N). Seta 2.25-4.0 mm long, 140-150 pm in di
ameter, 6 or 7 cells across (Figure 8G). Spores light
brown, hemispherical (Figure 10B), 35.0^42.5 pm in di
ameter, including marginally projecting lamellae; distal
face convex (Figure 10A, C, D), ornamented with up to
10 discontinuous lamellae, ± 2.5 pm long and 5.0-7.5 pm
apart, but interconnected with faint cross ridges, some
times forming incomplete areolae and frequently with pa
pilla-like or cross-like inclusions; proximal face (Figure
10E) with incomplete and not very distinct triradiate mark,
facets covered with low papillae and short irregular ridges,
up to 25 lamellae projecting around periphery. Elaters
light brown, 107.5-175.0 pm long, 7.5 pm wide in middle,
tapering to ends, 3-spiral or partly 2-spiral, covered with
fine, low papillae (Figure 10F).
Fossombronia straussiana is often mixed with other
Fossombronia species and grows on streambanks and in
seepage areas at several localities in the Northern Province
and Gauteng (Figure 4). More specimens of it, J. Braggins
91/191 and S.M. Perold 2654, were collected in 1991 in
Malawi on the Zomba Plateau, as well as at Nyika Nat.
Park, S.M. Perold 2663', thus it appears to be widespread.
Vanden Berghen (1965) reported F. husnotii (with hyaline
rhizoids) from the Congo Republic, Symoens 4329, and
from Tanzania ( = Tanganyika), Bryan 1036. The spores
of these specimens have much taller lamellae than those
F. straussiana and also appear to differ from those of F.
husnotii. In 1978 Vanden Berghen reported F. husnotii
from Shaba, Zaire (Malaisse 9039), without mentioning
the rhizoids, but presumably referred here because they
are hyaline. Scott & Pike (1988a) comment that the spores
of F. husnotii are extraordinarily variable and that more
research is required. Beside F. husnotii, which has mostly
hyaline rhizoids, Scott & Pike (1984) also described three
new Australian species, F. punctata, F. scrobiculata and
F. vermiculata, as having hyaline rhizoids on all or on
most plants.
Fossombronia straussiana has been named in honour
of Mrs Susan Strauss, owner of the Farm Geelhoutbosch,
where she has collected it a number of times, together
with other Fossombronia species.
This species is easily distinguished by its hyaline or
brownish rhizoids and dense, frilly leaves, its pseudope
rianth with lamellate, lateral outgrowths, by the spore or
namentation that usually has incomplete areolae with
Bothalia 27,1 (1997) 25
FIGURE 8.—Fossombronia straussiana. A-F, leaves; G, cross section o f seta; H, opened pseudoperianth; I, pseudoperianth from side; J, perigonial
bract and 2 antheridia; K, cross section of stem; L, detail of leaf margin; M, median leaf cells with oil bodies and chloroplasts; N, cells in
capsule wall. A-C, E, F, H, I, K, N, Strauss & Retief CHI3655, D, J, Perold & Koekemoer 3116a\ G, Strauss CHI3653', L, M, Strauss
CH I3651. Drawings by G. Condy. Scale bars: A-F, H, I, 500 pm; G, K, 100 pm; J, 250 pm; L-N, 50 pm.
26 Bothalia 27,1 (1997)
FIGURE 9.—Fossombronia straussiana. A, stem branching near apex; B, leaves crowded on stem; C, antheridium and archegonium in close proximity
between leaves; D, proximal leaf appendage curved around gametangia; E, close-up o f same; F, pseudoperianth with seta emerging from it.
A, F, Strauss 133', B, Perold &. Koekemoer 3124a', C-E, Perold & Koekemoer 3116a. A, x 7; B, x 10; C, x 66; D, x 15; E, x 106; F, x 23.
inclusions and by the usually finely papillose elaters. The
specimen, Scott 13, has a larger and more elaborate
pseudoperianth than those usually encountered in this spe
cies. Its spore ornamentation is very similar, however, and
the rhizoids are hyaline.
ACKNOWLEDGEMENTS
I wish to thank the referees of this article, Dr E.O.
Campbell and Prof. R. Stottler for their constructive com
ments as well as the kind people who have collected
specimens for me, particularly Dr H.F. Glen and Mrs
Susan Strauss, also my colleagues at the National Bo
tanical Institute, Ms Marinda Koekemoer and Mr Jac
ques van Rooy who helped me with fieldwork. Many
thanks to Dr Glen for translating the diagnoses into
Latin, Mrs J. Mulvenna and Ms D. Maree for typing
the manuscript, Ms G. Condy for the drawings and Mrs
A. Romanowski for developing and printing many pho
tographs.
FIGURE 10.—Fossombronia straussiana. Spores and elater. A, C, D, distal face; B, side view of distal face; E, proximal face; F, elater. A, B, E, Scott 13',
C, Strauss & Retief CH I3655', D, S.M. Perold3280', F, Perold & Koekemoer 3124a. A, C, x 727; B, x 1084; D, x 588; E, x 674; F, x 839.
Bothalia 27,1 (1997) 27
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SPECIMENS EXAMINED
Held at PRE, unless otherwise indicated. Bracketed
numbers after citation of collector’s name and collecting
number refer to the species described in the text in alpha
betical order, namely: F. gemmifera (1); F. glenii (2); and
F. straussiana (3).
Braggins 91/191 (3) (Malawi).
Glen 2134 (2) (paratype), 2146 (2) (holotype).
Koekemoer 976 (2).
Mogg CHI57 (2) BOL, PRE.
Perold S.M. 2017 (1); 3052 (2); 3280, 3281, 2654 (Malawi), 2663
(Malawi) (3). Perold & Koekemoer 3116a, 3124a (3); 3116b, 3124b,
3129 (1). Perold & Van Rooy 3555, 3559a (paratype), 3564, 3565 (1);
3568, 3569 (2).
Scott 13 (3) CH3697. Strauss 132 (1) (holotype), 133 (3) (holotype), 134
(3) (paratype), CH13651, CH13653, CH13654 (3). Strauss & R elief
CH13655 (3).
Wager 14 (3).
