FLORA OF SOUTHERN AFRI
BRYOPHYTA
Editor O. A. Leistner
Part I Mosses
Fascicle I
S p h agn aceae — G r i m m i aceae
by Robert E. Magill
Botanical Research Institute
Department of Agriculture and Fisheries
Republic of South Africa
FLORA OF SOUTHERN AFRICA
All contributions should be compiled in accordance with the Guide to Contributors to the Flora of Southern
Africa (Compiled by Ross, Leistner & De Winter) which is available from the Librarian, Botanical Research
Institute, Private Bag X101, Pretoria 0001.
Cryptogam volumes will in future not be numbered but will be known by the name of the group they cover
The number assigned to the volume on Characeae therefore becomes redundant.
Alien families are marked with an asterisk.
Published volumes and parts are shown in italics.
INTRODUCTORY VOLUMES
The genera of Southern African flowering plants
Vol. 1: Dicotyledons (Published 1975). Price: R1 1,00.’ Overseas: R14,00. Post free
Vol. 2: Monocotyledons (Published 1976). Price: R8,00. Overseas: R10, 00. Post free
Botanical exploration in Southern Africa
CRYPTOGAM VOLUMES
Charophyta (Published as Vol. 9 in 1978). Price: R4,25. Overseas: R5, 30. Post free
Bryophyta:
Part 1: Mosses: Fascicle 1: Sphagnaceae — Grimmiaceae ( Published 1981). Price: R24,33, GST R0,97,
Total R25,30. Overseas: R30,40. Post free
Fascicle 2: Gigaspermaceae — Bartramiaceae
Fascicle 3 : Erpodiaceae — Hookeriaceae
Fascicle 4: Fabroniaceae — Polytrichaceae
Part 2: Hepatophyta, Anthocerotophyta
Pteridophyta
FLOWERING PLANTS VOLUMES
Vol. 1 : Stangeriaceae, Zamiaceae, Podocarpaceae, Pinaceae*, Cupressaceae, Welwitschiaceae, Typhaceae,
Zosteraceae, Potamogetonaceae , Ruppiaceae, Zannichelliaceae, Najadaceae, Aponogetonaceae , Junca-
ginaceae, Alismataceae, Hydrocharitaceae (Published 1966). Price: Rl,75. Overseas: R2,20. Post free
Vol. 2: Poaceae
Vol. 3: Cyperaceae, Arecaceae, Araceae, Lemnaceae, Flagellariaceae
Vol. 4: Restionaceae, Mayacaceae, Xyridaceae, Eriocaulaceae, Commelinaceae, Pontederiaceae, Juncaceae
Vol. 5: Liliaceae, Agavaceae
Vol. 6: Haemodoraceae, Amaryllidaceae, Hypoxidaceae, Tecophilaeaceae, Velloziaceae, Dioscoreaceae
Vol. 7: Iridaceae
Vol. 8: Musaceae, Strelitziaceae, Zingiberaceae, Cannaceae*, Burmanniaceae, Orchidaceae
Vol. 9: Casuarinaceae*, Piperaceae, Salicaceae, Myricaceae, Fagaceae*, Ulmaceae, Moraceae, Cannabaceae*,
Urticaceae, Proteaceae
Vol. 10: Part 1 : Loranthaceae, Viscaceae (Published 1979). Price: R2,00. Overseas: R2,50. Post free
Santalaceae, Grubbiaceae, Opiliaceae, Olacaceae, Balanophoraceae, Aristolochiaceae, Rafflesiaceae,
Hydnoraceae, Polygonaceae, Chenopodiaceae, Amaranthaceae, Nyctaginaceae
Vol. 11: Phytolaccaceae, Aizoaceae, Mesembryanthemaceae
( cont. on back cover )
PRICE OF THIS FASCICLE: Local: R24,33, GST R0,97, Total R25,30.
Other countries: R30,40. Post free.
Printed by the Government Printer and obtainable from the Division of Agricultural Information,
Department of Agriculture and Fisheries, Private Bag X144, Pretoria 0001.
REPUBLIC OF SOUTH AFRICA
REPUBLIEK VAN SUID-AFRIKA
DEPARTMENT OF AGRICULTURE AND FISHERIES
DEPARTEMENT VAN LANDBOU EN VISSERYE
FLORA OF SOUTHERN AFRICA
BRYOPHYTA
PART 1, FASCICLE 1
ISBN 0 621 06951 5
G.P.-S.
Digitized by the Internet Archive
in 2016
https://archive.org/details/floraofsoutherna11unse
FLORA OF SOUTHERN AFRICA
which deals with the territories of
SOUTH AFRICA, TRANSKEI, LESOTHO, SWAZILAND, BOPHUTHATSWANA,
SOUTH WEST AFRICA/NAMIBIA, BOTSWANA AND VENDA
BRYOPHYTA
PART 1 MOSSES
Fascicle 1 Sphagnaceae — Grimmiaceae
by
Robert E. Magill
Edited by
O. A. Leistner
Editorial Committee: B. de Winter, D. J. B. Killick and O. A. Leistner
Botanical Research Institute,
Department of Agriculture and Fisheries
1981
CONTENTS
Page
New taxa and a new combination published in part 1, fascicle 1 vi
Geographical regions referred to in this fascicle vii
Introduction ix
Vegetation formations in the Flora area ix
Fynbos ix
Desert/Karoo x
Grassland/savanna x
Woodland x
Forest x
Alpine heath-grassland xi
Collecting, preservation, study and identification of the mosses of Southern
Africa xi
Collecting xi
Preservation xii
The moss packet xii
Study and identification xii
Characters used in keys and descriptions xiv
Illustrations xv
Glossary 1
Conspectus of classification 13
Literature cited 15
Provisional key to the families of fascicle 1 17
SPHAGNACEAE 23
ANDREAEACEAE 33
FISSIDENTACEAE 39
NANOBRYACEAE 69
ARCHIDIACEAE (by J. van Rooy) 71
DITRICHACEAE 83
SELIGERIACEAE 103
DICRANACEAE 105
CALYMPERACEAE 161
ENCALYPTACEAE 173
POTTIACEAE 177
BRYOBARTRAMIACEAE 269
GRIMMIACEAE 271
Index to fascicle 1 283
NEW TAXA AND A NEW COMBINATION PUBLISHED IN PART 1
FASCICLE 1
Acaulon recurvatum Magill, sp. nov., p. 199
Acaulon rufochaete Magill , sp. nov., p. 201
Andreaea bistratosa Magill, sp. nov., p. 37
Barbula microcalycina Magill, sp. nov., p. 241
Bruchia foveolata Magill, sp. nov., p. 108
Chorisodontium falcatum Magill, sp. nov., p. 154
Crossidium apiculatum Magill, sp. nov., p. 195
Crossidium spiralifolium Magill, sp. nov., p. 197
Fissidens capriviensis Magill, sp. nov., p. 45
Husnotiella plicata Magill, sp. nov., p. 222
Leucoperichaetium Magill, gen. nov., p. 271
Leucoperichaetium eremophilum Magill, sp. nov., p. 273
Pleuridium papillosum Magill, sp. nov., p. 88
Pottia namaquensis Magill, sp. nov., p. 206
Weisiopsis involuta Magill, sp. nov., p. 225
Desmatodon longipedunculatus (C. Mm//.) Magill, comb, nov., p. 210
Date of publication: October, 1981.
GEOGRAPHICAL REGIONS REFERRED TO IN THIS FASCICLE
B — Botswana
CC — central Cape
CE — eastern Cape
CN — northern Cape
CS — southern Cape
CNW — northwestern Cape
CSW — southwestern Cape
L — Lesotho
N — Natal
O — Orange Free State
S — Swaziland
SWA — South West Africa/Namibia
T — Transkei
TC — central Transvaal
TE — eastern Transvaal
TN — northern Transvaal
TS — southern Transvaal
TW — western Transvaal
Z — Zululand
vii
INTRODUCTION
The following text represents the first of four fascicles making up Part 1 of Volume
Bryophyta in the Flora of Southern Africa Cryptogam series. Part 1 of this volume will treat
the mosses, including Sphagnopsida, Andreaeopsida, Bryopsida and Polytrichopsida, Part 2
being reserved for the Liverworts and Hornworts. Fascicle 1 includes Sphagnopsida, Andreae-
opsida and roughly, the Acrocarpi-haplolepideae of the Bryopsida. The remaining three
fascicles will treat the Acrocarpi-diplolepideae and Pleurocarpi-diplolepideae of Bryopsida
and Polytrichopsida (see Conspectus of classification, p. 13).
The comparatively large temperate bryophyte flora of Southern Africa has a long and
interesting bibliographical history. Several Southern African taxa were treated in Hedwig’s
(1801) Species Muscorum Frondosorum and Hooker’s (1918-20) Musci Exotici and reference
has continuously been made to Southern African taxa up to and including Scott and Stone’s
(1976) Mosses of Southern Australia and Smith’s (1978) Moss Flora of Britain and Ireland.
Historical aspects of the bryological activity in Southern Africa have been dealt with in the
introduction of Sim’s (1926) Bryophyta of South Africa and, more recently, an account of
Southern African bryological collections and collectors was published by Magill (1980).
Further information will soon be made available by Gunn & Codd (1981).
Several excellent texts are available on the life history and morphology of mosses, e.g.
Flowers (1973), Parihar (1965) and Watson (1972); therefore only brief accounts of taxonomi-
cally important structures will be given in each fascicle.
Vegetation formations in the Flora area
Bryophytes are found within each of the six major vegetation formations encountered in
Southern Africa, namely fynbos, desert/karoo, grassland/savanna, woodland, forest and
alpine heath-grassland. The Southern African mosses are frequently more or less restricted,
at least at the species level, to one of these formations. Many taxa, however, show a broader
environmental tolerance and occur in two or more formations, or at least in transitional zones
between them. For example, species of Macrocoma are frequently collected in savanna,
woodland and forest formations (vide Magill & Vitt, 1981). Likewise, Fabronia is collected
over large areas of Southern Africa dominated by grasslands, savannas or woodlands.
A synopsis of the phytogeographical relationships of the Southern African mosses will
only be possible after completion of all four fascicles. A brief statement on each of the vege-
tation formations is given here as an aid to identification and as a provisional reference to
distribution patterns cited in the text.
Fynbos
Restricted primarily to the extreme southern tip of Africa, this vegetation formation con-
tains a unique flora of ericoid and restioid elements, spread across the mountains and coastal
lowlands of the southern and western Cape. Fynbos is not difficult to identify in the field, but
older collections can frequently only be tentatively placed within the fynbos formation. This is
primarily due to the rather discontinuous distribution of this formation, and the remnant
patches of forest or of karoo vegetation scattered throughout its area.
In addition it is also frequently difficult to determine whether or not the mosses are truly
part of the lowland fynbos. Mosses only rarely grow on the coarse quartzite sand inhabited
by fynbos elements, but are more commonly collected on bare rock outcrops or cliff- faces
within the formation.
Mosses are however, frequently collected in close association with mountain fynbos
elements that occur on richer soils of mountain slopes and in valleys.
IX
An indication of the mosses present in the fynbos was recently given by Schelpe (1970)
in a provisional checklist of the bryophytes of the Cape Peninsula. The dominant mosses in the
fynbos region are species of Sphagnaceae, Dicranaceae, Pottiaceae, Grimmiaceae and
Bryaceae. The family Wardiaceae is endemic to the fynbos.
Desert / Karoo
Phytogeographically the arid and semi-arid regions of the southern and western parts of
the Flora area are generally included in the Karoo-Namib region. This region includes areas
of bleak deserts, with large expanses devoid of vegetation in the west, but it extends eastward
into open dwarf-succulent shrublands. The vegetation becomes taller and woody on mountain
slopes and along temporary water courses. Bryophytes are practically unknown from the
drier desert areas of South West Africa/Namibia, and they are infrequently collected in the
shrublands of the Karoo.
The majority of the mosses from the arid and semi-arid parts of the western Cape and
southern South West Africa/Namibia are minute ephemerals of Ditrichaceae, Dicranaceae
and Pottiaceae, only rarely collected by the most astute collectors. Species of Pottiaceae and
Grimmiaceae are the most commonly encountered mosses in the Karoo of the central and
southern Cape. Generally disguised by their greyish or blackish colour when dry, these species
generally form extensive colonies on rocks or on soil below low shrubs. Although very in-
frequent, species of Funariaceae, Bryaceae and Fabroniaceae are occasionally encountered in
denser vegetation or in protected rock crevices.
Grassland/ savanna
The grasslands and savannas of the Zambezian Domain cover large expanses of the
northern and eastern Flora area, and extend northward to cover most of southcentral and
eastern Africa. Mosses are only occasionally found in the flatter areas throughout this region,
but become much more frequent as elevation increases or where the vegetation is denser,
especially around rock outcrops or on cliff faces. Members of the families Fissidentaceae,
Ditrichaceae, Dicranaceae and Pottiaceae are the most commonly encountered, although
Archidiaceae, Bryaceae, Orthotrichaceae, Fabroniaceae and Polytrichaceae are also present.
Composition of the moss flora differs in riverine situations or wooded areas along streams,
where Bryaceae, Funariaceae and occasionally members of Hypnales are found. Sphagnum
is frequently collected in the grassland/savanna formation in permanently wet habitats.
Woodland
Scattered throughout the grassland of the Zambezian Domain are numerous patches of
woodland. As interpreted here, the woodland formation encompasses a wide range of vege-
tation types, i.e. from dense savanna to open dry forests. The mosses of the woodlands include
many of the grassland taxa, but a higher percentage of the pleurocarpic families is characteristic
of this formation.
As the woodlands become denser, either in kloofs or on mountain slopes, species of
Erpodiaceae, Orthotrichaceae, Hedwigiaceae, Cryphaeaceae and Leskeaceae are frequently
encountered. Members of the Hypnales are more common in the thicker woodlands around
small mountain streams.
The wetter, high elevation woodlands of the escarpment in the eastern Transvaal and
Drakensberg of Natal are distinct bryologically and more closely related to the forest flora.
Forest
In Southern Africa, natural forest exists as large or small remnants scattered from Table
Mountain in the southwestern Cape, across coastal regions of the southern and eastern Cape,
and into Natal. The forests extend northward along the escarpment of the eastern Transvaal
and link up with the montane forests of eastern Africa.
x
The forest remnants vary in size from the larger Knysna and Amatola Forests in the Cape,
to the tiny kloof forests of Natal. In addition, several forest types are represented in Southern
Africa, i.e. montane, kloof-mist, lowland and coastal. Comparison of the bryophyte communi-
ties in each of these forest types is underway, but results are fragmentary at this time.
The forests support a large and diverse bryophyte flora. The most important difference
from the woodlands is the higher frequency of members of the Isobryales and Hookeriales.
Alpine heath-grassland
Confined chiefly to the Drakensberg of Lesotho and adjoining parts of Natal, Orange
Free State and eastern Cape, the alpine heath-grassland has a vegetation characterized by
alpine grasses and dwarf shrubs. The elevation, high precipitation and, to some extent, soil
types contribute to the unique status of this formation in Southern Africa. The extremely
interesting bryophyte flora of the alpine region is particularly rich in species of Dicranaceae,
Pottiaceae, Bryaceae, Bartramiaceae and Polytrichaceae. Numerous species with disjunct
distributions are also known from the area, i.e. Bryoerythrophyllum jamesonii, B. recurviro-
strum, Bryum albopulvinata, B. cyathiphyllum and Abietinella abietina.
Collecting, preservation, study and identification of the mosses of Southern Africa
Collecting
Most mosses can be collected at any time of the year, the one exception being the epheme-
rals of the drier regions. These tiny mosses are found only after the rainy season, e.g. Sep-
tember-October in Namaqualand. Different periods of fertility within the mosses will result
in the production of sporophytes on some species, during any part of the year.
Bryophytes are found throughout Southern Africa, from the wet forest in the east to the
arid deserts of the west and from the Cape fynbos to the Okavango Swamps in Botswana.
The mosses in dense, wet forests are large and easily collected ; consequently they are also
the best-known part of the moss flora. As precipitation decreases, the frequency, size and
prominence of the mosses also decrease. Changes in growth strategies and habitat preference
are also obvious.
The collecting of bryophytes is relatively simple when compared with other forms of
biological collecting. This is partly due to their small size, but also to the fact that practically
no field preparation is necessary. The most convenient collecting method employs small
paper bags, of c. 250 g-1 kg size, as specimen containers. Samples are generally removed from
the substrate by hand or with a small knife. A lOx hand lens is useful for initial observations.
Excess soil, humus or bark should be removed and a single collection placed in a paper bag.
Various methods are used for numbering specimens and recording the collecting locality
and substrate information. The most reliable procedure is to record all of this information on
the collecting bag. To save time, larger paper bags can be used to store all of the collecting
bags from a single locality and individual collection numbers may be added at a later time. The
substrate data must be noted as accurately as possible on each bag when the collection is
made.
It is very important to air-dry the specimens, as soon as possible, to avoid fungal growth.
For this reason plastic bags should be avoided when collecting or for storing field collec-
tions.
It must be emphasized that mosses represent an important part of any ecosystem not only
aesthetically but ecologically. Judicious collecting should always be undertaken with the
knowledge that mosses grow very slowly. Entire colonies may be eradicated as a result of
damage caused by sampling, or they may take many years to recover.
xi
Preservation
Mosses require minimal preparation before being identified or preserved as herbarium
specimens. Large, bulky specimens can be made pliable by wetting. After these specimens are
divided or folded to conform to packet size, they should be lightly pressed and dried between
sheets of blotting paper. This procedure should be avoided when dealing with very small
specimens. A fine soil sieve will remove excess soil, without disturbing delicate cushions or
specimens collected on crusts of soil.
The mosses are well suited to storage in a packet system, whether the packets are kept
separately or mounted on herbarium sheets. Vertical file packet systems are not only curatorial-
ly practical, but less expensive in both material and space. Containers used for the filing of
specimens can be as simple as a group of shoe boxes or as elaborate as specially made boxes in
broad-side filing cabinets (cf. Magill, 1980). Mounting bryophytes exposed on herbarium
sheets should be avoided.
Herbarium packets for bryophytes can be made to fit the size of an available box or
cabinet or designed to suit the collector. A useful standard packet can be made from an A4
piece of paper. The quality of the paper should be such that packets will last for many years.
The moss packet
(1) Place a sheet of A4 paper on a flat surface. Bring the lower edge of the paper
upward 100 mm for the first fold, square the sides and crease the fold;
(2) fold each side over 35 mm, square the lower edges, then crease each of the side
folds;
(3) fold the top downward to c. 7-8 mm above the bottom of the packet; square
the sides and crease the fold.
This will result in a 104x 138 mm packet suitable for most vertical file packet systems.
Collecting data for individual specimens (substrate, precise locality and altitude, sur-
rounding vegetation, date, collector and number, and name of the moss) should be written
on the outer flap or a label with this information can be glued to the flap. Some collectors
have the paper pre-printed so that when it is folded into a packet, the outer flap has a label
in place, ready for inscription of the information.
Study and Identification
Techniques used in the study and identification of mosses vary widely and undoubtedly
collectors, undertaking the identification of their own specimens, will modify the techniques
discussed here. The following method should be regarded as a starting point only.
A dissecting and a compound microscope, as well as the standard materials for preparing
microscope slides, are needed for the identification of mosses. Two pairs of fine-tipped forceps
and a sewing needle secured in a wooden handle are useful for the dissection of specimens.
A razor blade or scalpel is required to make transverse or longitudinal sections of stems,
leaves or capsules.
Before dissection begins, the specimen should be examined and notes should be made of
habit, colour, position and curvature of leaves and, if present, the position and structure of
the sporophyte. A single plant, or perhaps two or more, depending on size, should be wet in a
drop of water at one end of a slide; some scientists prefer to use a beaker of boiling water to
wet specimens. Either method will work. Position and curvature of the wet leaves should
again be noted.
Under the dissecting microscope, leaves from the upper stem are removed using both
pairs of forceps. One pair of forceps should be used to hold the stem in place and the other
either to scrape leaves off the stem (in a motion from the apex downward) or to pick off
xii
individual leaves (care should be taken to grasp the leaves at the insertion with the stem).
Perichaetial and perigonial leaves should not be mixed with the vegetative leaves, but segre-
gated with the sporophyte until later.
Free-hand sectioning of moss leaves or stems is never easy and beginners may make
numerous attempts before obtaining usable sections. Several methods of sectioning have been
published (Flowers, 1973; Scott & Stone, 1976; Zander, 1979a). Although all these methods
produce results, they tend to be rather hit or miss. In order to fully use all of the characters
available from leaf or stem sections, this Flora requires very precise sectioning, frequently in
specific areas of the leaves or stems.
In the method employed during preparation of the Flora a leaf is placed in a small drop
of water at the corner of a slide. Depending upon whether proximal or distal sections are
required, the apex or base of the leaf is held down with a pair of forceps or needle. Under the
highest magnification of the dissecting microscope, a single edge razor blade is employed in a
slow rocking motion over the leaf, across the corner of the slide. Careful positioning of the
blade will quickly produce precise, thin sections from various parts of the leaf. A similar
technique is used when stem and capsule sections are needed.
The detached leaves and sections are moved to another slide and placed into a drop of
water or mounting medium. A cover glass is then applied. As stated earlier the perichaetial
leaves and sporophyte should be kept separate from the vegetative leaves. When making a
permanent slide it is a good practice to have the two under separate cover slips, on the same
slide.
Wetting of the capsule frequently requires additional manipulation, i.e. boiling water or a
small drop of alcohol to wet the peristome and contents of the spore sac. In most cases the
capsule is too large to be mounted whole under a cover slip. The following procedure is
useful for obtaining the necessary parts.
The capsule is secured by forceps at a corner of the slide. The mouth and, if present,
associated peristome, is severed near the top of the urn. The resulting ring of the mouth is
then divided into 2-4 parts, being careful not to damage the peristome teeth. The urn should
then be sectioned lengthwise to produce a long narrow strip of the exothecial cells. The mouth
parts, urn section, perichaetial leaves and any other associated parts, are then transferred to a
drop of water or mounting medium and covered by a cover glass. Some of the mouth parts
should face up and others down so that both surfaces of the peristome teeth can be seen.
Spores are generally carried with parts of the capsule, however occasionally capsules must be
searched for a few spores. When permanent slides are made, the remainder of the capsule and
attached seta can be placed in the mounting medium alongside the cover slip for later reference.
Mosses are commonly identified from water-mounted preparations, however, a set of
permanent microscope preparations is continuously useful for comparison, especially if slides
are properly labelled with names and reference numbers. Although a variety of mounting
media is available, Hoyer’s medium is widely used in bryology becaues of its water solubility,
clearing quality and simple recipe.
To prepare Hoyer’s medium : (1) very slowly dissolve 30 g of gum arabic (preferably flake
type) in 50 ml of distilled water at room temperature. This is most easily accomplished in a
500 ml beaker on a magnetic stirrer, over 2-4 hours; (2) add 200 g of chloral hydrate, stir
until dissolved, then allow to stand till clear; (3) add 20 ml of glycerine and stir slowly; (4)
the solution should be kept in an air-tight container and small amounts decanted into a drop-
per bottle when needed.
Occasionally moss leaves shrink or contort when transferred directly from water to
Hoyer’s medium. Although the leaves generally respread quickly, problem groups, i.e. Funaria
and Rhodobryum, can be treated in a graduated series of glycerine/water when necessary.
The slides can be used immediately, but the medium will only harden over 1-2 weeks at
room temperature. The resultant semi-permanent slide will remain usable for many years;
xiii
ringing the cover slip with nail varnish will make the slide virtually permanent. All measure-
ments and observations in the Flora were made from specimen preparations mounted in
Hoyer’s medium.
Characters used in keys and descriptions of mosses
Plants. Characters describing the habit are somewhat subjective and generally represent an
impression of size, growth form and colour obtained from the specimens examined. The
substrate preference is taken from field observation and label information when available.
The size calegories, used in the descriptions, are roughly equated with the following list.
Some deviation may occur, especially where taxa span more than one category.
Plant size categories
Minute < 1mm high
Small <10 mm high or long
Medium 10-30 mm high or long
Large 30-60+ mm high or long
Robust >70 mm high or long but also referring to thick or full plants.
Stem. Transverse sections of stems are only infrequently required for identification of mosses
of this fascicle. One important exception is Sphagnum where the number of layers of the stem
hyalodermis is taxonomically important. In a few families, presence or absence of a central
strand is significant at the generic level. Ornamentation or unusual differentiation of the stem
surface cells are infrequent in taxa treated in this fascicle.
Leaves. Position and curvature of leaves are frequently important in identification and should
be observed in both dry and wet condition. Leaf size and shape, cited in the keys and descrip-
tions, are taken from leaves above the middle of the stem but below the apex, unless otherwise
stated. Basal stem leaves, perigonial and perichaetial leaves are frequently distinct from the
vegetative leaves; where appropriate, these leaves are also described.
Leaf length always indicates total length from base to apex. When necessary for identifi-
cation, lengths of parts such as subula or base are given separately, in addition to total leaf
length. Awns, however are not included in leaf length, but are recorded separately under the
costa.
Leaf shape refers to the lamina as a whole. Apical and basal shapes refer only to the
extreme tip or base of the leaf, respectively.
The leaf sections referred to in keys, descriptions and illustrations are cross sections
from the upper middle of the leaf, unless otherwise stated.
Costa. The length and strength, surface structure and anatomy of the costa provide many
important characters for the identification of the mosses in this fascicle. It is therefore generally
necessary to examine leaf sections when identifying the mosses of this fascicle.
Laminal cells. There are generally two, but occasionally many, differentiated cell regions in a
single leaf. Laminal cells and basal cells are the two types most frequently encountered and
are generally described. The shape of the laminal cells was taken from an area of the upper
third of the leaf, midway between the margin and centre-line. Basal cells are described from
a similar area in the lower part of the leaf, but above the cells at the insertion. Cell ornamenta-
tion and thickenings were observed in surface view and in transverse sections. Cell measure-
ments refer to the long axis unless otherwise stated.
Sporophyte. Shape and curvature of the capsule and seta are frequently important for identifi-
cation, and should be noted both wet and dry whenever possible. The shape of exothecial cells
and stomatal type and position are less important, but can be useful in identifying some
specimens.
XIV
Size, shape and ornamentation of the peristome teeth are important at all levels of classifi-
cation and should be studied when available. Presence or absence of a peristome is frequently
critical in identification. Fragile peristomes quickly break away giving the impression of a
gymnostomous capsule. Therefore capsule mouths should be routinely checked for remnants
of the old peristome.
The operculum and calyptra of mosses in this fascicle can generally be studied under the
dissecting microscope and need only rarely be included on slides. When the operculum or
calyptra are important for identification, they are usually large enough to be easily observed
under the dissecting microscope, e.g. Encalypta, Bryobartramia.
Spores are generally very uniform in size, shape and ornamentation. They are occa-
sionally very ornate ( Trematodon , Bruchia) or differ significantly in size or shape between
species ( Archidium , Fissidens). Spore measurements refer to maximum diameter unless other-
wise stated.
Illustrations
All illustrations were prepared by Mrs Rita Weber using pencil as the medium. A very
fine, soft lead (0,3 mm HB) was employed to give the sharp line necessary for reproduction,
but also to allow for the shading that makes pencil such an attractive medium for bryophyte
illustrations.
The habit drawings and their enlargements were prepared by the artist by direct observa-
tion of selected, generally wetted plants, using a dissecting microscope. All other parts were
drawn from photographs, taken by the author, using a photographic apparatus attached to a
compound microscope. The dissected specimens, used for photography, were mounted in
Hoyer’s medium.
XV
1
GLOSSARY
abaxial — the side away from the stem or
axis back or lower surface of a leaf
(opposed to adaxial) \ see dorsal.
acaulescent — essentially stemless or apparently
so.
acrocarpic — gametophyte producing sporo-
phyte at apex of a stem or main branch
(opposed to pleurocarpic). Acrocarpous
mosses generally grow erect in tufts (rather
than mats) and are sparsely branched,
acumen— a slender, tapering point,
acuminate — slenderly tapered. Fig. 1: 16.
acute — with angle of less than 90°. Fig. 1:15.
adaxial — toward the stem or axis, upper
surface of a leaf (opposed to abaxial );
see ventral.
alar cells — group of cells at the basal margins
of a leaf, at insertion; occasionally strongly
differentiated; e.g. Dicranoloma.
amplexicaul — clasping the stem,
anisophyllous — either bearing two distinct
types of leaves, e.g. Hypopterygium and
Racopilum; or having stem and branch
leaves obviously dissimilar, e.g. Porotham-
nium and Thuidium (opposed to isophyllous).
annulus — a ring of differentiated cells between
the mouth of the capsule and the oper-
culum, aiding in dehiscence,
antheridium — the male sex organ, a globose
to broadly cylindric, stalked structure
containing sperms (antherozoids); see
perigonium.
apiculate — abruptly short-pointed ( mucronate
is shorter; cuspidate is longer and stouter),
apiculus — a short, abrupt point,
appendiculate— with short, transverse projec-
tions (appendiculae); see also trabeculate.
appressed — closely applied, as in leaves to
the stems.
archegonium — the female sex organ, a flask-
shaped structure producing an egg; see
perichaetium.
arcuate — curved.
areolation — the cellular network of a leaf.
aristate — bristle-pointed, ending in an awn;
see hair-point.
articulate — with thickened joints, jointed,
ascending — pointing obliquely upward,
attenuate — slenderly tapering,
auriculate — with auricles.
auricle — a small, ear-like lobe at the basal
margins of a leaf.
autoicous — with archegonia and antheridia
in separate inflorescences on the same
plant; cladautoicous — with the male inflo-
rescence on a separate branch ; gonio-
autoicous — with the male inflorescence
bud-like and axillary on the same stem or
branch as the female inflorescence;
pseudautoicous — with dwarf male plants
epiphytic on the female; rhizautoicous —
with the male inflorescence on a very short
branch attached to the female stem by
rhizoids and appearing to be a separate
plant.
awn — a hair-point, usually formed by an
excurrent costa; e.g. piliferous leaves of
Tortula.
axil — the angle between leaf and stem,
axillary — in the leaf axils.
axillary hairs — uniseriate filaments found in
leaf axils, generally inconspicuous and
concealed by the leaf bases,
basal cells — cells of the lower £ to ^ of a leaf,
between margin and costa (opposed to
laminal cells).
basal membrane — in Haplolepideae a short
tube or cylinder of the peristome generally
supporting numerous teeth; e.g. Tortula.
In Diplolepideae a delicate but often well-
developed membrane at the base of the
inner peristome, commonly terminating in
segments and cilia.
beak — the prolonged apex of an operculum.
bordered — having margins differentiated from
the rest of the lamina in shape, size, colour,
or thickness of cells; see limbidium.
brood-body — reduced bud, leaf or branch
( propagulum ); e.g. Tortula pagorum; or
small, globose, ellipsoidal, or cylindrical to
filamentous, septate bodies ( gemmae ); e.g.
Barbula indica; serving in vegetative repro-
duction.
2
3
bloom— a powdery or waxy covering,
bulbiform — bul b-shaped .
caespitose — tufted, growing in cushions or
sods.
calyptra (pi. calyptrae) — a membranous hood
over the developing sporophyte, developed
from the upper part of the archegonium;
in true mosses ruptured near the base,
carried upward by the elongation of the
seta, and continuing growth to form a cap
over the capsule; see epigonium. Fig. 2:1.
campanulate — bell-shaped,
canaliculate — channeled,
cancellate — lattice-like,
cancellina (pi. cancellinae) — sharply defined,
large, empty, and usually hyaline basal
leaf cells of the Calymperaceae.
capsule — the spore case, or sporangium,
often differentiated into an upper spore-
bearing urn and a sterile basal portion,
the neck. Fig. 2:1.
cartilaginous — firm and tough,
catenulate — chain-like,
caulescent — having a stem,
central strand — a small group of elongate
cells forming a central axis of some stems,
usually thin-walled or coloured in trans-
verse section. Fig. 2: 6.
cernuous — nodding or drooping,
chlorocyst — small, elongate, green cells form-
ing a network surrounding the large,
hyaline cells ( leucocysts ) of a Sphagnum
leaf; also applied to small, green cells
enclosed by layers of hyaline cells in the
leaves of Leucobryum.
chlorophyllose — green, containing chlorophyll,
cilia — delicate, thread-like structures alter-
nating with the segments of the inner
peristome of many mosses; also applied to
hair-like appendages fringing leaves or
calyptrae. Fig. 2: 3.
ciliate — fringed with hairy appendages,
circinate — curved in a circle,
cladautoicous — see autoicous.
clavate — club-shaped.
cleistocarpic — referring to indehiscent cap-
sules lacking an operculum or valves and
hence opening irregularly (opposed to
stegocarpic)-, e.g. cleistocarpous capsule of
Bruchia and Archidium.
cochleariform — rounded and deeply concave,
shaped like the bowl of a spoon,
collenchymatous — with cell walls more strong-
ly thickened at the corners; thickenings
called trigones.
columella — the central sterile tissue in a
capsule around which the spores develop,
commissure — margin; in leaves of Sphagnum,
the junction between the adjacent walls of
hyalocyst and chlorocysts; pores along the
commissures are thus arranged along the
lateral margins of hyalocysts.
comose — hairy; but used by bryologists to
refer to larger and more crowded leaves
forming tufts or comae at the stem tips;
e.g. Campylopus.
complanate — flattened together or compres-
sed in 1 plane; e.g. Fissidens.
complicate — folded lengthwise,
complicate-carinate — sharply folded along a
keel.
conduplicate — strongly folded along the mid-
dle; e.g. Eustichia.
constricted— abruptly narrowed,
contorted — bent into irregular curves, irre-
gularly twisted.
Fig. 1. — Leaf shapes (1-14): 1. setaceous above an ovate base, 2. linear, 3. triangular, 4. lanceolate, 5. obo-
vate, 6. oblong, 7. spathulate, 8. ovate, 9. orbicular, 10. ligulate, 11. lingulate, 12. elliptical, 13. falcate, 14. subulate
above an oblong, clasping base. Leaf apices and costal lengths (15-20): 15. acute with percurrent costa, 16.
acuminate with percurrent costa, 17. obtuse with excurrent costa, 18. rounded with costa ending below apex,
19. mucronate apex, 20. cuspidate apex. Marginal curvature of laminae seen in cross section (21-25) : 21. incurved
(left lamina only), 22. recurved (right lamina only), 23. plane (complete cross section). 24. involute (left lamina
only), 25. revolute (right lamina only). Laminal cell shapes (26^ 33): 26. rounded, 27. quadrate, 28. rectangular,
29. hexagonal, 30. linear, 31. fusiform, 32. rhombic, 33. oblong-hexagonal (rhomboidal). Laminal cell ornamen-
tation seen in superficial view (above) and cross section (below) (34-42): 34. flat and smooth, 35. slightly bulging
and smooth, 36. mammillose, 37. sharply mammillose, 38. papillae low and blunt, 39. papillae C-shaped, 40.
papillae bifid (spinose), 41. papillae crown-shaped, 42. cells prorate.
4
convolute — rolled together and forming a
sheath.
cordate — heart-shaped .
cortex — differentiated outer layers of a stem
or branch, occasionally surrounding a
central strand.
cortical — referring to the outer cells of stem
or branches, frequently differentiated into
inner and outer layers. Fig. 2: 6.
corticolous — growing on bark.
cosmopolitan — occurring in all major floristic
zones of the world.
costa — nerve or midrib of a leaf, sometimes
double.
costate— with a costa.
crenate— with rounded teeth.
crenulate — with minute, rounded teeth.
cribrose — finely perforated.
crisped (crispate) — wavy (like crisp bacon);
often used more loosely to mean variously
curled, twisted and contorted.
cryptopore — immersed stoma, with the guard
cells sunken below the level of the exo-
thecial cells and often ± covered by them
(opposed to phaneropore).
cucullate — hooded or hood-shaped; a cucul-
late calyptra is conic and split up 1 side,
resembling a monk’s hood; also used to
describe leaves concave at the tips (cf.
mitrate ); e.g. Tortula, Campylopus.
cupulate — cup-shaped, rounded and swollen;
e.g. capsule of Gigaspermaceae.
cushion — a closely-packed aggregate of plants
with stems ± erect, but somewhat radiating
at edges ; e.g. Grimmia.
cuspidate — ending abruptly in a stout, rigid
point; cf. apiculate. Fig. 1 : 20.
cyathiform — cup-shaped, slightly wider at
top; e.g. capsules of Phasconica.
cygneous — curved downward like a swan’s
neck.
cymbiform — concave and broadly boat-
shaped.
deciduous — -falling off, caducous.
decumbent — lying prostrate but with ascend-
ing tips.
decurrent — with margins extending down the
stem below the leaf insertion as ridges or
narrow wings.
dehiscent— splitting open, referring to cap-
sules opening by valves (or by an oper-
culum)', e.g. Andreaea.
dendroid— branched above a trunk-like stipe
and resembling a tree; e.g. Porothamnium.
dentate — with sharp teeth directed outward;
see tooth.
denticulate — finely dentate.
deoperculate — referring to a capsule after the
operculum has fallen.
depauperate — poorly developed.
dextrorse — twisted to the right (opposed to
sinistrorse).
diaphanous — transparent.
dimorphic — of two forms.
dioicous — with archegonia and antheridia on
separate plants, dioecious.
distal — away from the base or point of
attachment (opposed to proximal).
distichous — in 2 opposite rows; e.g. Fissi-
dens, Distichium.
divisural line — the median line of a peristome
that is usually zig-zag, sometimes fur-
rowed.
dorsal — the back, lower, or abaxial surface
of a leaf; the upper surface of a prostrate
stem; or the outer surface of a peristome
tooth (opposed to ventral).
dorsal lamina — in Fissidens the portion of the
leaf blade at the back of the costa, opposite
the sheathing base (cf. vaginant laminae).
dorsal plates — the outer surface of exostome
teeth.
dorsiventral — flattened, with distinct upper
and lower surfaces.
echinate— roughened by blunt, spiny projec-
tions.
ecostate — without a costa.
elliptical— essentially oblong with convex
sides or ends. Fig. 1:12.
emarginate — broad at apex and shallowly
notched ; e.g. Tortula.
5
emergent — partially exposed, referring to
capsules only partly exposed beyond tips
of perichaetial leaves.
endemic— limited to a single country or
floristic area.
endostome — the inner peristome (usually con-
sisting of segments, frequently alternating
with cilia, above basal membrane); e.g.
Diplolepideae. Fig. 2: 3.
entire — without teeth, ± smooth, generally
referring to leaf margins.
epigonium — protective envelope of the de-
veloping sporophyte before it ruptures into
two parts, a vaginula that encircles the
base of the seta and foot of the sporo-
phyte, and a calyptra that covers the cap-
sule. Bryobartramia has a persistent epigo-
nium.
epiphragm — a circular membrane formed by
the expanded tip of the columella, partially
closing the mouth of the deoperculate
capsule; e.g. Polytrichum and Funaria.
epiphyllous — growing on leaves of higher
plants.
equitant — straddling, referring to the con-
duplicate and strongly sheathing leaf bases
of Fissidens.
erect — upright.
erect-spreading — patent, spreading at an
angle of 45° or less; cf. spreading.
erose — irregularly notched or ragged, as
though gnawed.
excurrent — referring to a costa that extends
beyond the apex. Fig. 1: 17.
exostome — the outer peristome, consisting of
teeth; e.g. Diplolepideae. Fig. 2: 3.
exothecium — the outermost layer of the
capsule wall, consisting of exothecial cells.
exserted — projecting and exposed, referring
to capsules projecting above tips of peri-
chaetial leaves.
falcate — curved like the blade of a sickle.
Fig. 1: 13.
falcate-secund — strongly curved and turned to
one side.
fasciculate — bunched together, in bundles or
fascicles; e.g. branches of Sphagnum.
fibrillose— with fine, fibre-like wall thicken-
ings (fibrils); applied to hyaline cells of
Sphagnum where the fibrils may be spiral or
annular.
filiform — slender and elongate, filamentous,
thread-like.
fimbriate — f ringed .
flaccid — soft, limp, flabby.
flagellum — a slender branch; referring to
minute, axillary brood-branches and some-
times to long, slender, tapering stems
(flagellate).
flexuose — slightly and irregularly bent, twist-
ed, or wavy.
foot — the basal absorbing organ of the
sporophyte.
foveolate — marked with small pits; e.g. spores
of Bruchia foveolata.
frond — the branched part of a dendroid
or frondose moss.
frondose — closely and regularly branched in
one plane (pinnate), resembling a fern
frond; e.g. Thuidium.
fruit — the capsule or more loosely the sporo-
phyte.
fugacious — vanishing or readily falling away.
fusiform — narrow and tapered at both ends,
spindle-shaped. Fig. 1:31.
gametophore — the leafy phase of the gamete-
bearing generation, produced from buds
on the protonema.
gametophyte — the dominant, sexual genera-
tion; in mosses, the green, leafy plant.
gemma (pi. gemmae) — a small, globose,
elliptic, or cylindric body of a few cells
serving in vegetative reproduction; see also
brood-body and propagulum.
gemmiferous — bearing gemmae.
gemmate — bud-like.
glabrous — smooth; neither papillose, rough
nor hairy.
glaucous — with a whitish, greyish, or bluish
bloom.
globose — spherical.
granulose (granulate) — roughened with mi-
nute, blunt projections; minutely grainy.
6
gregarious — growing close together but not
in tufts or mats.
gonioautoicous — see autoicous.
guide cells — large, empty, rather thin-walled
cells in a median row as viewed in the cross
section of the costa of many mosses.
Fig. 2: 4-5.
gymnostomous — lacking a peristome.
gynoecium — the female inflorescence ; see
perichaetium.
habit — the aspect or general appearance of a
plant.
habitat — local environment.
hair-point — see awn and aristate, e.g. Grim-
mia laevigata.
heterogeneous — dissimilar (opposed to homo-
geneous).
heteroicous— with several forms of inflore-
scence on the same plant (or on various
plants of the same species); also called
polyoicous.
heteromallous — pointing in all directions (op-
posed to homomallous).
hispid — with short, stiff hairs ; bristly.
hoary — whitish or greyish, usually referring
to plants with leaves long hair-pointed;
e.g. Grimmia.
homogeneous — uniform (opposed to hetero-
geneous).
homomallous — pointing the same way (op-
posed to heteromallous).
hyaline — colourless and transparent.
hyalocyst — large, empty, colourless water-
storage cell, surrounded by a network of
green chlorocysts; see also leucocysf, e.g.
Sphagnum leaves.
hyalodermis — a cortex of large, empty, colour-
less cells ( hyalocysts ) ; e.g. Sphagnum.
hydroids — tracheid-like conductive cells in
the axial strand of many mosses, some-
times also in the costa but not usually in
contact with the central strand.
hymenostomaceous — peristome lacking but
capsule mouth closed by an ephemeral
membrane developed from the top of the
columella.
hypophysis — a swollen neck at the base of the
capsule (also called apophysis),
imbricate — closely appressed and overlap-
ping.
immersed — completely covered; referring to
capsules exceeded by the tips of the peri-
chaetial leaves.
incised — cut into sharp divisions separated
by narrow sinuses,
incrassate — thickened.
incurved — curved upward and inward, ap-
plied to leaf tips and margins (opposed to
recurved ); cf. erect. Fig. 1 : 21.
inflexed — bent upward and inward, applied
to leaf margins (opposed to reflexed).
inflorescence — a cluster of sex organs and the
leaves or bracts surrounding them; see
also perichaetium and perigonium.
innovation — a new shoot; a branch formed
after the maturity of sex organs, usually
at the base of an inflorescence,
insertion — the place of attachment of a leaf,
branch, or peristome.
internal cylinder — central axis of Sphagnum
stem, inside hyalodermis; cf. central strand.
involute — rolled upward and inward, applied
to leaf margins (opposed to revolute). Fig.
1 : 24.
irregular — asymmetric,
isodiametric — about as broad as long (includ-
ing square, rounded, or hexagonal),
isophyllous — stem and branch leaves similar
(opposed to anisophyllous).
julaceous— smoothly cylindric, like a catkin,
referring to stems or branches with crowded
and imbricate leaves.
juxtacostal— adjacent to costa.
keeled — sharply folded along the middle,
like the keel of a boat.
lacerate — deeply and irregularly slashed or
torn.
laciniate — dissected into fine, deep, irregular
divisions (laciniae); ciliate-fringed.
lamella (pi. lamellae) — green ridges or plates
on the costa or lamina of some moss leaves.
lamina (pi. laminae) — the leaf blade (as
distinguished from the costa).
7
lamina! cells — cells of the upper £ of a leaf,
between the margin and costa (opposed
to basal cells).
lanceolate— lance-shaped, narrow and tapered
from the base (narrowly ovate-acuminate).
Fig. 1 : 4.
lateral— at the side (opposed to terminal).
lax — soft or loose; usually referring to large,
thin-walled cells, but also the nature and
spacing of leaves on stem, or stems in
tufts.
lenticular (lentiform) — doubly convex, lens-
shaped ; cf. gemmae of Oedipodiella.
leptoid — conductive cell, somewhat similar
to sieve tubes in form and function, found
in the axial strand of stems and setae of
Polytrichaceae and in setae of many other
mosses.
leptome — a phloem-like tissue consisting of
leptoids and parenchymatous cells.
leucocyst — large, empty, hyaline leaf cells of
Sphagnum and Leucobryum; see also
hyalocyst.
lid — operculum.
ligulate — strap-shaped ; narrow, moderately
long, with parallel sides. Fig. 1 : 10.
limbate — with a limbidium; e.g. Fissidens.
limbidium— a border of elongate, incrassate,
marginal leaf cells that are distinct from
other laminal cells; e.g. Fissidens.
linear — very narrow, elongate, with nearly
parallel sides (narrower than ligulate).
Fig. 1 : 2 & 30.
Ungulate — tongue-shaped; oblong with a
slightly broadened apex. Fig. 1:11.
lumen — the cell cavity.
lunate — (lunulate) — crescent-shaped.
mammilla (pi. mammillae) — a bulging pro-
tuberance.
mammillose (mammillate) — convex with a
blunt central projection. Fig. 1 : 36-37.
mat — a densely interwoven, horizontal form
of growth; e.g. Br achy the cium, Hypnum.
micrometre — one-thousandth of a millimetre,
micron (abbreviated — pm).
midrib — a mid-vein or single costa.
mitrate (mitriform) — conic and undivided or
equally lobed at base, similar to a bishop’s
mitre, referring to calyptrae, cf. cucullate;
e.g. Grimmia, Ptychomitrium.
monomorphic — of a single form,
monoicous — with antheridia and archegonia
on the same plant, including autoicous,
paroicous, polyoicous and synoicous (op-
posed to dioicous).
monopodial — main stem of unlimited growth;
sex organs produced on lateral branches
(opposed to sympodial).
mucro — a short, abrupt point,
mucronate — ending abruptly in a short point,
usually caused by a shortly excurrent
costa ( apiculate is somewhat longer; cuspi-
date is even longer and stouter). Fig. 1:19.
muticous — without a point or awn.
nanandrous — producing dwarf males.
neck — the sterile basal portion of a capsule,
sometimes considerably differentiated, (op-
posed to urn) cf. hypophysis', e.g. Trema-
todon.
nerve — costa; midrib.
nodose — with knob-like thickenings, i.e. cell
wall thickenings of Grimmiaceae.
nodulose — minutely knobbed,
oblate — wider than long,
oblong — rectangular. Fig. 1 : 6 & 28.
obscure — indistinct.
obsolete — scarcely evident, almost lacking;
usually referring to the costa.
obtuse — broadly pointed, more than 90° —
used by some authors to mean blunt or
rounded. Fig. 1:17.
opaque — not transparent or translucent.
operculum— the lid covering the mouth of a
moss capsule, falling at maturity to release
the spores. Fig. 2:1.
orbicular — nearly circular. Fig. 1 : 9.
oval — shortly elliptical. Fig. 1: 12.
ovate — egg-shaped (with the basal portion
broader than the apical portion). Fig. 1 : 8.
panduriform — shaped like the body of a
violin (obovate with a rounded sinus on
either side).
8
Colyptra
f— Operculum
Ventral surface
Cells
Guide ceils
Dorsal steroid
band
Dorsal Surface
cells
4
Ventral surface
cells
Ventra I stereid
band
Guide cells
Dorsal Stereid
band
Dorsal surface
cells
Outer cortex
Central Strand
Inner cortex
Fig. 2. — 1. Moss gametophyte and sporophyte. 2-3. Peristome types: 2. single (haplolepidous), 3. double
(diplolepidous). 4-5. Costal cross sections. 6. Stem cross section.
9
papilla (pi. papillae)— a minute protuberance
of various forms. Fig. 1 : 38-41.
papillose — roughened by 1 to many minute
protuberances. Fig. 1: 38-41.
paraphyllia— small, green, filiform, lanceo-
late, leaf-like or sometimes branched
scales ; often produced on stems or branches
of pleurocarpous mosses; e.g. Thuidium.
paraphyses — hyaline or yellowish, septate
hairs mingled with the antheridia (and
often with archegonia).
parenchyma — a tissue of relatively undif-
ferentiated cells, usually thin-walled and
isodiametric.
parenchymatous — composed of ± short cells
joined end to end (opposed to prosen-
chymatous).
paroicous — with antheridia and archegonia
in the same inflorescence but not mixed,
the antheridia in the axils of uppermost
and perichaetial leaves.
patent — erect-spreading.
pellucid — clear, translucent or transparent.
pendent — hanging.
pendulous — somewhat drooping, inclined
more than horizontally; with stems and
branches hanging, a growth form common
in forests.
percurrent — referring to a costa that extends
to the apex; cf. excurrent. Fig. 1: 15-16.
perichaetium (pi. perichaetia) — the female
inflorescence, consisting of leaves and the
enclosed archegonia.
perigonium (pi. perigonia) — the male inflo-
rescence, a cluster of leaves and the
enclosed antheridia.
peristome — a single (haplolepidous) or double
(diplolepidous) circular structure inside
the mouth of the capsule ; in diplolepidous
mosses the outer peristome ( exostome )
consists of teeth, the inner peristome
( endostome ) consists of segments, some-
times alternating with cilia, and often
rising from a basal membrane. Fig. 2: 1-3.
phaneropore — superficial stoma, with the
guard cells at the same level as other
exothecial cells and not sunken in chambers
(opposed to cryptopore).
phyllodioicous — nanandrous; with dwarf male
plants resting on leaves or tomentum of
the larger female plants.
phyllotaxy — the arrangement of leaves on
a stem.
piliferous — hair-pointed; e.g. awn.
pilose — with long hairs; usually referring to
leaves with long-excurrent costa.
pinnate — with numerous, spreading branches
on 2 sides of the axis and thus resembling
a feather.
pitted — with small openings or pores through
the walls of adjoining cells, sometimes
marked by conspicuous depressions in
thick-walled cells (also called porose).
plane — flat, referring to leaf margins. Fig.
1: 23.
pleurocarpic — producing the sporophytes
laterally from a perichaetial bud or a
short, specialized branch rather than at the
stem tips (opposed to acrocarpic)', pleuro-
carpous mosses are usually prostrate,
creeping and freely branched and grow in
mats rather than tufts.
plicate — folded in longitudinal pleats (plicae).
plumose — closely and regularly pinnate;
feathery.
polymorphic — of many forms, variable.
polyoicous — with antheridia and archegonia
in separate inflorescences mixed together
on the same plants (also called heteroicous).
pore — small opening or ‘pit’ in the walls of
some cells, sometimes conspicuous as
depressions in thickened walls between
adjacent cells; in Sphagnum the pores are
particularly large and conspicuous in the
exposed walls of the leucocysts of branch
leaves and also in outer stem cells; porose
cell walls may be minutely or quite obvious-
ly perforated.
porose — pitted ; see pore.
primary stem — the main stem, often creeping
or rhizome-like with reduced or scale-like
leaves.
processes — see segment.
procumbent — spreading, prostrate.
proliferous — continuing growth by the pro-
duction of a new stem (or innovation).
10
propagulum (pi. propagula) — reduced bud,
branch, or leaf serving in vegetative repro-
duction; see also brood-body.
prorate — with leaf cell ends projecting above
the surface of the lamina giving a papillose
appearance or with papillae (mammillae)
borne at one or both ends of a cell (also
called scindulose); e.g. Philonotis. Fig.
1: 42.
prosenchyma — a tissue made up of elongate
cells with tapered ends,
prosenchymatous — with narrow, elongate cells
overlapping at the ends (opposed to
parenchymatous).
prostome (preperistome) — a rudimentary
structure outside and usually adhering to
the peristome teeth,
prostrate — creeping.
protonema (pi. protonemata) — green, branch-
ed filaments produced on germination of
spores and giving rise to a leafy gameto-
phyte; in Sphagnum and Andreaea, the
protonema is ± thallose rather than fila-
mentous ; secondary protonemata are some-
times produced from broken leaves or
stems of mosses.
proximal — near the base or point of attach-
ment (opposed to distal),
pseudautoicous — see autoicous.
pseudoparaphyllia — small, unistratose, lan-
ceolate or filiform structures resembling
paraphyllia, often found in pleurocarpous
mosses, in small numbers around branch
primordia.
pseudopodium — an elongation of the game-
tophytic axis below the sporophyte in
Sphagnum and Andreaea, serving the
function of a seta; also applied to a similar
extension of a stem tip bearing clusters of
gemmae.
pseudopore — small spots somewhat resem-
bling pores in leucocysts of Sphagnum
branch leaves; consisting of fibril rings not
enclosing a perforation.
pulvinate — cushion-like.
punctate — dotted, usually referring to spore
markings.
pyriform — pear-shaped .
quadrate — square. Fig. 1 : 27.
radicles — rhizoids; slender, non-chlorophyl-
lose threads attaching the plants to the
substrate, often covering stems, occasion-
ally growing from leaf tips; the radicles of
mosses usually have oblique cross-walls;
cf. rhizoids.
radiculose — covered with rhizoids.
recurved — curved downward and inward ;
referring to leaf margins or tip, marginal
teeth, or peristome teeth (opposed to
incurved). Fig. 1 : 22.
reflexed — bent down and inward, generally
referring to leaf margins (opposed to
inflexed).
regular — symmetric.
reniform — kidney-shaped.
repent — creeping.
resorption — disappearance or erosion of parts
of cell walls (resorbed); e.g. Sphagnum
leaves.
resorption furrow — a furrow produced by the
resorption of the outer cell walls of the
elongated marginal cells in leaves of some
species of Sphagnum; seen in cross section
as a C-shaped marginal cell.
reticulate — in a network.
retort cells — cells with a pore at the upper
end, terminating a short, projecting neck,
found in the hyalodermis of many species
of Sphagnum.
retuse — with a slight indentation of a broad
apex.
revoluble — rolling away, referring to an
annulus which falls in a broken ring.
revolute — rolled downward and backward,
referring to leaf margins (opposed to
involute). Fig. 1 : 25.
rhizautoicous — see autoicous.
rhizoids — simple or branched, septate fila-
ments, dead at maturity, anchoring the
plant and sometimes ± covering the
stem; cf. radicles. Fig. 2:1.
rhizome — a slender, horizontal, subterranean
stem (also called subterranean stolon);
e.g. Rhodobryum.
rhombic — diamond-shaped. Fig. 1 : 32.
rhomboidal — longer than hexagonal, oblong-
hexagonal. Fig. 1 : 33.
11
rosette — a compact cluster of leaves encirc-
ling the stem,
rostrate — beaked .
rugose — transversely wrinkled or undulate,
rugulose — with weak, transverse wrinkles,
saxicolous — growing on rocks,
scabrous — rough.
scalpelliform — asymmetric and resembling
the blade of scalpel.
scarious — dry and thin, scale-like or mem-
branous.
secund — turned to one side,
segment— a tooth-like division of the inner
peristome ( endostome ), also called processes
Fig. 2: 3.
septate — divided by cell walls; having parti-
tions.
seriate — in rows, e.g. uni-, bi-, multiseriate;
applied to ranks of leaves on a stem,
rows of cells or papillae, cf. stratose.
serrate — saw-toothed; with marginal teeth
pointing forward,
serrulate — minutely serrate,
sessile — without a stalk or seta,
seta (pi. setae) — stalk supporting the capsule.
Fig. 2:1.
setaceous — bristle-like. Fig. 1:1.
sheathing — surrounding and clasping the stem,
the base of the seta or the capsule,
shoulder — the area where a leaf base is
abruptly narrowed to the upper lamina,
sinistrorse — twisted to the left (opposed to
{dextrorse).
sinuate (sinuose) — wavy,
sinuolate — minutely wavy,
spathulate — tapering proximally from a
broad, rounded apex (similar to Ungulate
but more markedly narrowed from a
broad apex). Fig. 1 : 7.
spiculate — covered with fine points,
spinose — referring either to leaf margins or
costae with sharply pointed teeth, e.g.
Syrrhopodon gomesi'r, or very high, sharp
leaf cell papillae, e.g. Triquetrella tristicha.
Fig. 1 : 40.
sporangium — the spore-sac of a capsule or,
more loosely, the entire capsule.
spores — minute, mostly spherical, nearly
always unicellular bodies produced in the
capsule as a result of reduction division,
giving rise on germination to protonemata
Fig. 2: 1.
sporophyte — the spore-bearing generation;
produced by the fertilization of an egg;
remaining attached to the gametophyte
and partially dependent on it, typically
consisting of foot, seta and capsule.
spreading — an angle of 45° or more; cf. wide-
spreading, erect-spreading, squarrose.
squarrose — spreading at right angles,
squarrose-recurved — spreading at right angles,
with the tips curved downward,
stegocarpic— referring to capsules with oper-
culum differentiated (opposed to cleisto-
carpic ); e.g. stegocarpous capsules of
Ditrichum and Barbula.
stellate — star-shaped.
stereids — slender, elongate, thick-walled fibre-
like cells found in groups (stereid bands)
in the costa of some mosses. Fig. 2: 4-5.
stipe — the unbranched part of an erect stem
of dendroid or frondose mosses,
stoloniform — referring to slender, elongate,
spreading branches or stems, with smaller
bract-like leaves.
stoma (pi. stomata) — opening in the capsule
wall, usually in the neck, surrounded by
guard cells.
strangulate — deeply constricted, as below
the mouth of a capsule,
stratose — in distinct layers; uni-, bi-, multi-
stratose; e.g. bistratose leaves have laminal
cells in two layers, cf. seriate.
striate— marked with fine, longitudinal ridges
(striae).
striolate — finely ridged,
strict — straight and rigid.
strumose — with a goitre-like swelling (or
struma) at the base, applied to some
capsules.
strumulose — with a small or indistinct swelling
at one side of the capsule base.
substereid — almost with the characters of a
stereid, walls not as strongly thickened.
subula — a long, slender point.
12
subulate — slenderly long-acuminate. Fig. 1:14.
sulcate — strongly plicate, with deep longi-
tudinal folds.
superficial cells — outer cells of the costa in
surface view; cf. surface cells.
surface cells — outer cells of the costa as seen
in cross section; i.e. superficial cells in cross
section. Fig. 2: 4-5.
sympodial — main stem of determinate growth;
sex organs produced at apex; main stem
frequently branching from below apex
(opposed to monopodial).
synoicous— with antheridia and archegonia
in the same inflorescence,
systylious — with the operculum remaining
attached to the tip of the columella after
dehiscence.
teeth — divisions of a single peristome, Fig.
2: 2; or of the exostome of a moss with a
double peristome. Fig. 2: 3; divisions of
the endostome are called segments; see
tooth and dentate.
teniola — intramarginal border of linear cells
in the leaves; e.g. Calymperes.
terete — rounded in cross section,
terminal — at the apex (opposed to lateral ).
terricolous— growing on soil,
tessellated — checkered, in a pattern of
squares, applied to the basal membrane of
the peristome of Tortula.
tetrad — a group of 4 developing or rarely
mature spores.
tetrad scar— a triangular to almost circular
feature occasionally obvious on the distal
face of some spores, resulting from the
contact between spores in a tetrad during
cell wall deposition; e.g. Bruchia and
Trematodon.
thallose — consisting of a flat plate of tissue,
rather than filaments; see protonema.
tomentose — woolly, densely radiculose, cov-
ered by a tomentum of rhizoids.
tomentum — woolly hairs or radicles.
tooth — a division of the peristome, Fig. 2:
2-3; see teeth', also applied to irregularities
or projections at the margins of leaves; see
dentate.
tortuose — irregularly bent or twisted.
trabeculate — with projecting cross-bars (tra-
beculae) at the back of some peristome
teeth; see appendiculate.
tristichous — in 3 rows; e.g. leaves of Tristi-
chium and Triquetrella.
truncate — abruptly cut off or squared off at
the apex.
tubulose — tube-like; usually referring to leaves
with strongly incurved margins; e.g. Cam-
pylopus.
tuft — growth form with stems erect, radiating;
small cushions; caespitose growth form,
turf — growth form with stems erect, parallel,
often in extensive colonies,
turgid — plump or swollen,
uncinate — hooked .
undulate— wavy.
urceolate — urn-shaped; applied to capsules
constricted below a wide mouth and
abruptly narrowed at the base,
urn — spore-bearing portion of a capsule
(opposed to neck).
vaginant laminae — in Fissidens, the equitant
laminae at base of the leaf that clasp the
stem and the base of the leaf above it;
cf. dorsal lamina.
vaginula — the ring or sheath enveloping the
base of the seta, derived from the base of
the archegonium and remaining after the
separation of the calyptra; see epigonium.
valve — one of the segments into which a
capsule of Andreaea separates on
dehiscence.
venter — the swollen basal portion of an
archegonium, containing the egg.
ventral — the upper or adaxial surface of a
leaf; the lower surface of a prostrate
stem; or the inner surface of a peristome
tooth (opposed to dorsal).
ventricose — bulging on one side below (like
a stomach).
vermiculate — with minute worm-like orna-
mentation; e.g. spores of Bruchia ecklo-
niana.
verruculate — irregularly roughened,
weft — a loosely interwoven, often ascending
growth form; e.g. Thuidium.
whorled — arranged in a ring or circle,
widespreading — spreading at a wide angle,
but less than 90°; cf. spreading.
13
CONSPECTUS OF CLASSIFICATION
The classification system used here is a linear model that inadequately expresses the rela-
tionships evident within the Musci.
The system is basically that of Fleischer (1902-22) and Brotherus (1924-25), modified to
accommodate some more recent concepts and it represents an intermediate stance between
the extremes of several published ordinal classifications.
It is obvious from recent research (vide Koponen, 1977; Edwards, 1979) that the classi-
fication of mosses needs realignment to conform with present concepts. This Flora is not
considered the proper venue for such alterations.
DIVISION BRYOPHYTA
Fascicle 1 :
CLASS SPHAGNOPSIDA
ORDER SPHAGNALES
Family Sphagnaceae
CLASS ANDREAEOPSIDA
ORDER ANDREAEALES
Family Andreaeaceae
CLASS BRYOPSIDA
ORDER DICRANALES
Family Fissidentaceae
Nanobryaceae
Archidiaceae
Ditrichaceae
Seligeriaceae
Dicranaceae
ORDER POTTIALES
Family Calymperaceae
Encalyptaceae
Pottiaceae
Bryobartramiaceae
Grimmiaceae
Fascicle 2 :
ORDER FUNARIALES
Family Gigaspermaceae
Ephemeraceae
Funariaceae
Splachnaceae
ORDER BRYALES
Family Bryaceae
Mniaceae
Eustichiaceae
Rhizogonaiceae
Aulacomniaceae
Bartramiaceae
Fascicle 3:
ORDER ORTHOTRICHALES
Family Erpodiaceae
Rhachitheciaceae
Ptychomitriaceae
Orthotrichaceae
Rhabdoweisiaceae
Racopilaceae
ORDER ISOBRYALES
Family Fontinalaceae
Wardiaceae
Hedwigiaceae
Cryphaeaceae
Leucodontaceae
Prionodontaceae
Trachypodaceae
Pterobryaceae
Meteoriaceae
Phyllogoniaceae
Neckeraceae
Lembophyllaceae
14
ORDER HOOKERIALES
Family Hookeriaceae
Fascicle 4 :
ORDER THUIDIALES
Family Fabroniaceae
Leskeaceae
Thuidiaceae
ORDER HYPNOBRYALES
Family Amblystegiaceae
Brachytheciaceae
Entodontaceae
Plagiotheciaceae
Hypnaceae
Hylocomiaceae
CLASS POLYTRICHOPSIDA
ORDER POLYTRICHALES
Family Polytrichaceae
15
LITERATURE CITED IN FASCICLE 1
Bartram, E. B., 1939. Mosses of the Philip-
pines. Philipp. J. Sci. 68: 1-437.
Bizot, M., 1967. Quelques mosses africaines
et americaines. Bull. Soc. bot. Fr. 114:
423-427.
Bizot, M., Friis, I., Lewinsky, J. & Poes, T.,
1978. East African bryophytes IV. Danish
collections. Lindbergia 4: 259-284.
Bizot, M. & Kilbertus, G., 1979. Les Cam-
pylopus africains subg. Lucidus Biz. et Kilb.
Revue bryol. lichen. 45: 61-95.
Bizot, M. & Poes, T., 1979. East African
bryophytes. III. Acta bot. hung. 25:
223-261.
Bizot, M., Poes, T. & Sharp, A. J., 1979.
Results of a bryogeographical expedition to
east Africa in 1968, II. J. Hattori bot. Lab.
45: 145-165.
Brotherus, V. F., 1924-1925. Musci. In
Engler & Prantl (eds), Die naturlichen Pflan-
zenfamilien. Edn 2, Vol. 10-11. Leipzig.
Buck, W. R., 1979. A re-evaluation of the
Bruchiaceae with the description of a new
genus. Brittonia 31: 469-473.
Crosby, M. R. & Magill, R. E., 1977. A
dictionary of mosses. Missouri Botanical
Garden, St Louis.
Dixon, H. N., 1920. New and interesting
South African mosses. Trans. R. Soc. S.
Afr. 8: 179-224.
Dixon, H. N., 1922. Some new genera of
mosses. J. Bot., Lond. 60: 101-110.
Dixon, H. N., 1932. African mosses collected
by O. A. H</>eg. K. norske Vidensk. Selsk.
Skr. 1932, 4: 3-27.
Dixon, H. N. & Gepp, A., 1923. Rehmann’s
South African mosses. Kew Bull. 6:
193-238.
Dixon, H. N. & Wager, W. A., 1930. New
and noteworthy mosses from South Africa.
Trans. R. Soc. S. Afr. 18: 247-261.
Eddy, A., 1977. Sphagnales of tropical Asia.
Bull. Br. Mus. nat. Hist. 5: 359-445.
Edwards, S. R., 1979. Taxonomic implica-
tions of cell patterns in Haplolepidous moss
peristomes. In: Bryophyte Systematics.
Systematics Association, special volume
14. London: Academic Press.
Edwards, S. R., 1980. A revision of west
tropical African Calymperaceae. I. Intro-
duction and Calymperes. J. Bryol. 1 1 :
49-93.
Fleischer, M., 1902-1922. Die Musci der
Flora von Buitenzorg. Vols 1-4. Flora von
Buitenzorg, Leiden.
Flowers, S., 1973. Mosses: Utah and the west.
Provo: Brigham Young University Press.
Frahm, J.-P., 1974. Zur Unterscheidung und
Verbreitung von Campylopus introflexus
(Hedw.) Brid. und C. polytrichoides De Not.
Revue bryol. lichen. 40: 33-44.
Frahm, J.-P., 1976. Taxonomische Notizen
zur Gattung Campylopus II. Revue bryol.
lichen. 42: 603-616.
Gangulee, H. C., 1969-1972. Mosses of
eastern India and adjacent regions. Vol. 1,
fasc. 1-3, Calcutta.
Gradstein, S. R. & Sipman, H. J. M., 1978.
Taxonomy and world distribution of
Campylopus introflexus and C. pilifer
(=C. polytrichoides ): a new synthesis.
Bryologist 81: 114-121.
Grout, A. J., 1928-1840. Moss Flora of
North America. Vols 1-3, Newfane,
Vermont.
Gunn, M. D. & Codd, L. E., 1981. Botanical
exploration in Southern Africa. In press.
Hedwig, J., 1801. Species Muscorum frondo-
sorum. Lipsiae.
Hooker, W. J., 1818-1820. Musci exotici con-
taining figures and descriptions of new or
little-known foreign mosses and other cryp-
togamic subjects. Vols 1& 2, London.
Iwatsuki, Z. & Pursell, R. A., 1980. Axil-
lary hyaline nodules in Fissidens (Fissiden-
taceae). J. Hattori bot. Lab. 48: 329-335.
16
Koponen, T., 1977. Modern taxonomical
methods and the classification of mosses.
Congres International de Bryologie. Bry-
ophytorum Bibliotheca 13: 443-481.
Magill, R. E., 1980. Musci austro-africani II.
Bryophyte collections in southern Africa
and southern African type specimens in the
National Herbarium, Pretoria. Bothalia 13:
127-133.
Magill, R. E. & Schelpe, E. A., 1979. The
bryophytes of southern Africa: An anno-
tated checklist. Mem. bot. Surv. S. Afr. 43 :
1-38.
Magill, R. E. & Vitt, D. H., 1981. The phy-
togeography and ecology of Macrocoma
(Orthotrichaceae, Musci) in Africa. Both-
alia 14: 463-466.
Mitten, W., 1860. On some new species of
Musci and Hepaticae in the herbarium of
Sir W. J. Hooker, collected in tropical
Africa, chiefly by the late Dr Vogel and Mr
Barter. Trans. Linn. Soc. Lond. 23: 51-58.
Muller, C., 1849-1851. Synopsis Muscorum
frondosorum. Vols 1-2, Berlin.
Parihar, N. S., 1967. An introduction to Em-
bryophyta. Vol. 1. Bryophyta. Allahabad:
Central Book Depot.
Portier de la Varde, R., 1927. Mouses de
1’Oubangui. Archs Bot. Mem. 1, 3: 1-153.
Richards, P. W. & Clear, I. D., 1967. Notes
on African mosses. III. Campylopus and
Microcampylopus. Trans. Br. bryol. Soc. 5:
305-315.
Saito, K., 1975. A monograph of Japanese
Pottiaceae (Musci). J. Hattori bot. Lab.
39: 373-537.
Schelpe, E. A., 1969. New records of
Southern Hemisphere bryophyta for South
Africa. Jl S. Afr. Bot. 35: 109-112.
Schelpe, E. A., 1970. A provisional checklist
of the bryophytes of the Cape Peninsula.
Contr. Bolus Herb. 2: 49-70.
Schf- pe, E. A., 1979. Corrections and addi-
tions to the Musci in Sim’s Bryophyta of
South Africa. Trans. R. Soc. S. Afr. 44:
113-122.
Scott, G. A. M. & Stone, I. G., 1976. The
mosses of southern Australia. London:
Academic Press.
Sim, T. R., 1926. The bryophyta of South
Africa. Trans. R. Soc. S. Afr. 15: 1-475.
Smith, A. J. E., 1978. The moss flora of Britain
and Ireland. Cambridge: Cambridge Uni-
versity Press.
Snider, J. A., 1975. A revision of the genus
Archidium (Musci). J. Hattori bot. Lab.
39: 105-201.
Stone, I. G., 1971. The sporophyte of Tortula
pagorum (Milde) De Not. Trans. Br. bryol.
Soc. 6: 270-277.
Stone, I. G., 1976. Alaticosta, a new subgenus
of Acaulon in Australia. J. Bryol. 9:
213-227.
Theriot, I., 1929. Musci. In H. Schinz (ed.),
Beitrage zur Kenntnis der afrikanischen
Flora (XXXIV), neue Folge. Vjschr. naturf.
Ges. Zurich 74: 99-100.
Townsend, C. C., 1978. Notes on mosses
from Ceylon and India, 1-5. J. Bryol. 10:
125-137.
Wager, H. A., 1914. Some new South African
mosses. Trans. R. Soc. S. Afr. 4: 1-8.
Watson, E. V., 1964. Structure and life of
bryophytes. London: Hutchinson University
Library.
Zander, R. H., 1972. Revision of the genus
Leptodontium (Musci) in the new world.
Bryologist 75: 213-280.
Zander, R. H., 1977. The tribe Pleuro-
weisieae (Pottiaceae, Musci) in Middle
America. Bryologist 80: 232-269.
Zander, R. H., 1978. New combinations in
Didymodon (Musci) and a key to the taxa in
North America north of Mexico. Phyto-
logia 41: 11-32.
Zander, R. H., 1979. Notes on Barbula and
Pseudocrossidium (Bryopsida) in North
America and an annotated key to the taxa.
Phytologia 44: 177-214.
Zander, R. H., 1979a. Techniques for study
of Pottiaceae. Taxon 28 : 643-644.
17
PROVISIONAL KEY TO THE FAMILIES OF FASCICLE 1
The following key is provided to allow access to the families treated in this fascicle. When
necessary, couplets are also incorporated into the key, that refer to taxa which will only be
treated in fascicle 2, 3 or 4. A key to all families represented in the Flora will be included in
fascicle 4.
1 Plants pleurocarpic or of pleurocarpic habit or appearing acrocarpic but branches
arising from a creeping stem or rhizome (Fascicles 2-4)
1 Plants acrocarpic or of acrocarpic habit:
2 Plants cleistocarpic, generally minute to small :
3 Plants acaulescent or nearly so :
4 Protonema persistent, plants scattered on protonema
EPHEMERACEAE (Fascicle 2)
4 Protonema quickly disappearing or persistent only at base of individual plants:
5 Epigonium rupturing early; calyptra very small POTTIACEAE (p. 177)
5 Epigonium persistent or calyptra swollen, completely covering sporophyte:
6 Leaves ligulate to Ungulate, 0,3-0, 5 mm long; laminal cells small, papillose
BRY OBARTRAMIACEAE (p. 269)
6 Leaves larger, narrowly spathulate, 3-4 mm long; laminal cells lax, smooth
FUNARIACEAE (Fascicle 2)
3 Plants with well defined stem:
7 Spores very large, 50-290 //m; capsules globose, exothecial cells with large
brown spots at maturity ARCHIDIACEAE (p. 71)
7 Spores smaller, 10-15 ^m; capsules globose to pyriform, exothecial cells variously
coloured:
8 Capsules oval to globose, immersed DITRICHACEAE (p. 83)
8 Capsules elliptical to cylindrical or pyriform, emergent to exserted:
9 Capsule pyriform, neck well defined; spores ornate. .DICRANACEAE (p. 105)
9 Capsule without neck; spores weakly papillose POTTIACEAE (p. 177)
2 Plants stegocarpic or capsules dehiscent through longitudinal valves:
10 Plants minute; leaves secund; capsules pendent, operculate, exserted laterally
through leaves on arcuate seta DITRICHACEAE (p. 83)
10 Plants larger; seta occasionally arcuate but not exserting capsule laterally:
1 1 Plants reddish black to dark green occasionally greyish because of long hyaline
awns; growing mostly on rock or stony soil:
12 Laminal cells papillose or mammillose; leaves always costate:
13 Laminal cells papillose POTTIACEAE (p. 177)
13 Laminal cells mammillose:
14 Lamina completely bistratose above base
PTYCHOMITRIACEAE (Fascicle 3)
14 Lamina unistratose or only bistratose juxtacostally or marginally
POTTIACEAE (p. 177)
12 Laminal cells smooth or, if papillose, leaves ecostate:
15 Leaves incurled, crisped to contorted dry, muticose: plants dark green,
glossy PTYCHOMITRIACEAE (Fascicle 3)
18
15 Leaves appressed when dry, little contorted:
16 Plants blackish to reddish black, dull; leaves muticose; laminal cell walls
straight, mostly irregularly thickened; capsule dehiscent through 4
longitudinal valves ANDREAEACEAE (p. 33)
16 Plants blackish green to greyish, occasionally glossy; leaves mostly
piliferous or with hyaline tips; laminal cell walls sinuate or nodose,
occasionally straight; capsules operculate . . . . GRIMMIACEAE (p. 271)
11 Plants green to light green or yellowish green, occasionally whitish:
17 Laminal cells differentiated into small chlorocysts and large leucocysts:
18 Leaves unistratose; leucocysts surrounded by network of chlorocysts;
leucocysts frequently fibrillose, generally with numerous pores on exposed
walls SPHAGNACEAE (p. 23)
18 Leaves multistratose; chlorocysts enclosed by 2-8 layers of leucocysts :
19 Leaves plane above, recurved to squarrose dry; chlorocysts in distal
section triangular, in two rows; capsules erect, symmetric; peristome
teeth 8 CALYMPERACEAE (p. 161)
19 Leaves concave above, appressed to contorted dry; chlorocysts in distal
section rhombic, in single row; capsule curved, asymmetric; peristome
teeth 16 DICRANACEAE (p. 105)
17 Laminal cells not differentiated into chlorocysts and leucocysts:
20 Leaves with filaments or lamellae on ventral surface:
21 Plants medium to large or robust POLYTRICHACEAE (Fascicle 4)
21 Plants small to minute POTTIACEAE (p. 177)
20 Leaves without filaments or lamellae on ventral surface:
22 Leaves distichous or tristichous:
23 Leaves tristichous:
24 Leaves appressed wet or dry; laminal cells smooth
DITRICHACEAE (p. 83)
24 Leaves spreading wet; laminal cells spinose-papillose
POTTIACEAE (p. 177)
23 Leaves distichous and complanate:
25 Leaves with vaginant laminae clasping stem and a well-developed
dorsal lamina FISSIDENTACEAE (p. 39)
25 Leaves without dorsal lamina:
26 Laminal cells papillose (Fascicle 2)
26 Laminal cells smooth:
27 Plants minute; leaves elliptical-subulate, to 1 mm long
NANOBRYACEAE (p. 69)
27 Plants larger; leaves oblong-subulate, 3-4 mm long
DITRICHACEAE (p. 83)
22 Leaves in many rows around the stem, not distinctly distichous or tristi-
chous, occasionally complanate:
28 Laminal cells smooth:
29 Laminal cell walls sinuate to nodose GRIMMIACEAE (p. 271)
19
29 Laminal cell walls straight, occasionally strongly incrassate and
pitted :
30 Alar cells differentiated, enlarged and hyaline to coloured:
31 Costa broad, occupying y to | width of leaf base
DICRANACEAE (p. 105)
31 Costa narrow, occupying less than £ width of leaf base:
32 Leaves with narrow border of elongated, hyaline cells
DICRANACEAE (p. 105)
32 Leaves without differentiated marginal cells
SELIGERIACEAE (p. 103)
30 Leaves with alar and basal cells occasionally somewhat larger or
hyaline, but alar cells not strongly differentiated:
33 Plants covered with glaucous bloom; growing in shallow
caves or rock recesses DITRICHACEAE (p. 83)
33 Plants without glaucous bloom:
34 Stems julaceous, leaves clasping stem wet or dry, oval to
orbicular DICRANACEAE (p. 105)
34 Leaves variously spreading wet:
35 Plants small to minute; leaves small, oval to ovate,
piliferous, capsules gymnostomous, immersed or
exserted, cupulate with very broad mouth; or leaves
broad elliptical-apiculate, plants producing discoid
gemmae at apex
GIGASPERMACEAE/FUNARIACEAE (Fascicle 2)
35 Plants mostly larger; capsules not cupulate; gemmae, if
present, on rhizoids or axillary on stem:
36 Peristome double, occasionally single or absent; leaves
generally broad, acute to shortly acuminate, frequently
crowded at apex, //"leaves narrower, margins strongly
toothed to doubly serrate; laminal cells generally lax
(Fascicle 2)
36 Peristome single, rarely absent; leaves generally nar-
rower; laminal cells smaller, quadrate to rectangular,
mostly incrassate:
37 Plants corticolous or saxicolous, forming small tufts:
38 Plants corticolous; capsules cylindrical, emergent;
peristome single or double; calyptra large,
covering capsule ORTHOTRICHACEAE (Fascicle 3)
38 Plants saxicolous; capsules exserted; peristome
single; calyptra small, mitrate
PTYCHOMITRIACEAE (Fascicle 3)
37 Plants terricolous or on rock at waterfalls or seeps,
gregarious or forming large turfs; capsules exserted,
peristome single ; calyptra small, cucullate :
39 Leaf apices obtuse to rounded; or, if acute, leaves
widest above middle and margins crenulate to
serrate POTTIACEAE (p. 177)
20
39 Leaf apices acute to acuminate or subulate:
40 Leaf margins bistratose above base :
41 Leaves lanceolate to ovate-acuminate, incurved
dry POTTIACEAE (p. 177)
41 Leaves oval to oblong, subulate, flexuose to
contorted dry DICRANACEAE (p. 105)
40 Leaf margins unistratose:
42 Leaf margins narrowly revolute above base to
near apex or reflexed below, plane above:
43 Costa in section with well-defined ventral
stereid band; capsule ± asymmetrical,
generally curved and furrowed dry; peri-
stome teeth lanceolate, c'eft to near base
DITRICHACEAE (p. 83)
43 Costa in section without ventral stereid band,
ventral cells in 2-3 layers, similar to guide
cells, incrassate; capsule erect, cylindrical,
smooth; peristome weakly twisted, teeth
linear POTTIACEAE (p. 177)
42 Leaf margins plane to erect or rarely weakly
inflexed :
44 Capsules long-cylindrical; peristome teeth
divided to base, papillose throughout; stem
in section with outer cortical cells in 2-3
rows, stereids or strongly incrassate; cells
of leaf subula narrowly rectangular to
linear or elliptical and strongly incrassate
DITRICHACEAE (p. 83)
44 Capsules short-elliptica or cylindrical
above well-defined neck; peristome teeth
perforated or cleft to near middle, longi-
tudinally striate or barred, papillose above;
stem in section with outer cortical cells in
1-2 rows, incrassate; cells of leaf subula
generally broader, rectangular to short
rectangular or infrequently quadrate
DICRANACEAE (p. 105)
28 Laminal cells papillose, mammillose or prorate:
45 Laminal cells prorate:
46 Plants small; leaves oval to oblong, subulate; capsule cylindrical
with well-defined neck DICRANACEAE (p. 105)
46 Plants medium to large; leaves ovate-acuminate to lanceolate, if
subulate, plants large; capsule globose to short-cylindrical;
peristome double BARTRAMIACEAE (Fascicle 2)
45 Laminal cells mammillose or papillose:
47 Laminal cells mammillose:
48 Laminal cells distinctly mammillose dorsally and ventrally:
49 Plants growing on soil; leaves ligulate, 1-2 mm long, lamina
unistratose DICRANACEAE (p. 105)
21
49 Plants growing on rock ; leaves ovate-acuminate, 4-5 mm long,
lamina bistratose PTYCHOMITRIACEAE (Fascicle 3)
48 Laminal cells mammillose ventrally, flat to ± bulging dorsally:
50 Lamina or margins completely or in part bistratose:
51 Margins bistratose, laminae unistratose; leaves gemmi-
ferous apically; basal leaf cells differentiated as cancel-
linae CALYMPERACEAE (p. 161)
51 Lamina bistratose; gemmae, if present, axillary; basal
leaf cells not strongly differentiated :
52 Upper lamina bistratose juxtacostally
POTTIACEAE (p. 177)
52 Upper lamina completely bistratose
PTYCHOMITRIACEAE (Fascicle 3)
50 Lamina and margins unistratose:
53 Leaves acuminate above obovate base; apex acute, cucul-
late ; juxtacostal cells weakly papillose dorsally
DICRANACEAE (p. 105)
53 Leaves elliptical to obovate; apex broadly acute to
rounded ; juxtacostal cells smooth dorsally
POTTIACEAE (p. 177)
47 Laminal cells papillose (see also Orthotrichaceae, Fascicle 3) :
54 Papillae massive, crown-shaped, multifid :
55 Leaves bordered by 2-6 rows of elongate, hyaline cells;
alar cells distinct DICRANACEAE (p. 105)
55 Leaves not bordered by elongate, hyaline cells; alar cells
not differentiated POTTIACEAE (p. 177)
54 Papillae smaller, mostly low and blunt or C- to O-shaped,
occasionally spinose or bifid :
56 Margins narrowly bordered by elongate, hyaline cells or
bistratose and generally dentate :
57 Basal leaf cells differentiated as cancellinae
CALYMPERACEAE (p. 161)
57 Basal leaf cells not differentiated as cancellinae :
58 Costa in section with ventral stereid band exposed,
ventral surface cells not differentiated
DICRANACEAE (p. 105)
58 Costa in section with differentiated ventral surface
cells above ventral stereid band
CALYMPERACEAE (p. 161)
56 Margins not bordered, rarely bistratose :
59 Leaves broad, elliptical to Ungulate or spathulate ; laminal
cells with several C-shaped papillae over lumen :
60 Calyptra very large, cylindrical-rostrate, completely
covering capsule; peristome absent or teeth short-
lanceolate, fragile; costa ending below apex to
mucronate ENCALYPTACEAE (p. 173)
22
60 Calyptra smaller, cucullate ; peristome teeth divided into
32 filaments above basal membrane; costa apiculate
to long excurrent POTTIACEAE (p. 177)
59 Leaves narrower, mostly broadest below mid-leaf, ovate
to lanceolate or ligulate; papillae mostly low and blunt,
occasionally spinose or C-shaped; calyptra small,
cucullate POTTIACEAE (p. 177)
23
SPHAGNACEAE
Plants generally robust, forming large hummocks, pale green to yellow-green or occasionally
pinkish to reddish; terricolous or saxicolous, aquatic to semi-aquatic. Stems erect, sparsely
divided; hyalodermis transparent, internal cylinder brownish, pinkish or greenish; in section
central cells of internal cylinder thin-walled, becoming smaller, thickened and coloured toward
margin, cells of hyalodermis lax, in 1-4 rows. Fascicles of branches spirally arranged around
stem, becoming crowded at stem tip; branches monomorphic or dimorphic; internal cylinder
frequently coloured or green; in section hyalodermis 1 layer thick. Leaves generally differen-
tiated, ecostate; stem leaves smaller, distant; triangular to oblong or Ungulate; apex acute to
rounded, serrulate to lacerate; margin plane to erect, frequently bordered by 1-8 rows of
narrow cells; branch leaves numerous, frequently crowded; oval or ovate to lanceolate; apex
acute to obtuse, serrate to dentate or truncate with numerous teeth ; margins erect to incurved
above, bordered by 1-3 rows of narrow cells to near apex. Ar eolation of two types: leucocysts
large, inflated, hyaline cells, generally with numerous fibrils and pores (or pseudopores),
although pores and/or fibrils occasionally absent; chlorocysts much smaller, green, forming
network around leucocysts; in section chlorocysts triangular to trapezoid or oval to rectan-
gular, totally enclosed or exposed on one or both surfaces, leucocysts convex or strongly
bulging on one or both surfaces.
Dioicous or monoicous. Perigonia and perichaetia on short lateral branches ; perichaetial
leaves conspicuous, larger, convolute. Sporophyte elevated above perichaetial leaves at maturity
by pseudopodia, seta not differentiated; capsule operculate, gymnostomous, globose at
maturity, urceolate to oblong-cylindrical after dehiscence; operculum convex; calyptra
membranous, hyaline; spores expelled through explosive dehiscence, tetrahedal, relatively
large, weakly ornamented.
The family contains just over 200 species in a single genus, Sphagnum. The family is economically by far the
most important of the mosses. In addition to its ecological importance in bog formation, Sphagnum has a great
many uses in horticulture, agriculture, medicine and the chemical industry. In areas where Sphagnum grows in
abundance, it has also been used as a fuel and in the production of charcoal.
SPHAGNUM
Sphagnum L., Sp. PI. 2: 1106 (1753); Hedw., Spec. Muse. 27 (1801); Warnst., Sphag. Univ.
39(1911); Sim, Bryo. S. Afr. 129(1926); Garside in J1 S. Afr. Bot. 15: 59(1949); Eddy in Bull.
Br. Mus. nat. Hist. 5 (7): 371 (1977). Lectotype species: S. palustre L., vide Britt, in Britt., FI.
Bermuda 431 (1918).
With characters of the family.
The genus is found throughout the world in association with wet or damp habitats. In Southern Africa,
true bogs, in the northern sense, are not formed and Sphagnum is confined to shaded mountain seeps, stream
banks, small pools or open swampy areas.
1 Branch leaf margin (in section) with resorption furrow 3. S. strictum subsp. pappeanum
1 Branch leaf margin (in section) without resorption furrow:
2 Stem leaf leucocysts without fibrils:
3 Stem leaf apex very broad, lacerate; with distinct areas of narrower cells on either side of lower leaf
1.5. fimbriatum
3 Stem leaf apex acute to obtuse, frequently toothed, not lacerate; lower leaf without areas of narrower
cells:
4 Plants pink; chlorocysts in section broadly exposed ventrally 2. S. obtusiusculum
4 Plants pale green; chlorocysts in section totally enclosed or occasionally exposed dorsally
7. S. pycnocladulum
24
Sphagnaceae
2 Stem leaf leucocysts with fibrils, strong or weak:
5 Hyalodermis of branches, (in surface view) fibrillose 4. S.perichaetiale
5 Hyalodermis of branches efibrillose:
6 Stem hyalodermis 1 cell thick or outer cells of internal cylinder not thickened and hyalodermis
not well defined:
7 Branch leaves broadly oval, margins generally incurved; stem leaves approaching size of branch
leaves; chlorocysts in section equally exposed on dorsal and ventral surfaces 5. S. africanum
7 Branch leaves ovate, margins erect; stem leaves generally smaller; chlorocysts in section with
wider exposure on dorsal surface 6. S. truncatum
6 Stem hyalodermis 2-3 cells thick:
8 Pores of branch leaf leucocysts few; plants pink 2. S. obtusiusculum
8 Pores of branch leaf leucocysts numerous ; plants greenish :
9 Stem leaf leucocysts efibrillose or fibrils irregular, not well defined 7. S. pycnocladulum
9 Stem leaf leucocysts with regular, well defined fibrils 8. S', capense
1. Sphagnum fimbriatum Wils. in Hook.,
Crypt. Bot. Antarct. Voyage 92 (1845);
Warnst., Sphag. Univ. 53 (1911); Garside in
J1 S. Afr. Bot. 15: 68 (1949); Smith, Moss FI.
Brit. Irel. 44 (1978). Type: Falkland Island,
Hooker s.n. (BM).
Plants long, slender, pale green; terri-
colous. Stems 80-120 mm long; internal cylin-
der green to brownish green, hyalodermis
Map 1 . — • Sphagnum truncatum
x Sphagnum fimbriatum
efibrillose, pores few; in section inner cells of
internal cylinder thin-walled, becoming smal-
ler and thickened toward margin, hyalodermis
fragile, 2 cells thick. Fascicles of 4-5 dimor-
phic branches: 2-3 spreading, 1-2 pendent;
internal cylinder greenish, hyalodermis efibril-
lose, pores present on most cells. Stem leaves
spathulate, 1 ,0-1 ,4 mm long; apex entensive-
ly resorbed, rounded, lacerate; margins plane,
border weak, 2-3 rows of narrow cells
restricted to base ; lower portion of leaf with
obvious region of narrow cells on either side
of the midline; leucocysts efibrillose, pores
absent, dorsal and ventral surfaces extensively
resorbed in upper portion of leaf, frequently
leaving only a network of chlorocysts. Branch
leaves ovate-acuminate, 1,0-1, 8 mm long;
apex acute, with 1-2 teeth; margins incurved
above, border of 2-3 rows of narrow cells
reaching upper leaf; leucocysts regularly
fibrillose, pores numerous along commissures;
in section chlorocysts trapezoid with wider
exposure on ventral surface.
Perichaetia lateral ; leaves broad, spathu-
late, concave, apex truncate, lacerate. Pseudo-
podia 5-10 mm long; capsules short-cylindri-
cal, 2 mm long, red-brown; operculum con-
vex; spores 20-25 pm, with distinct tetrad
scar, smooth. Fig. 3: 1-9.
Fig. 3. — Sphagnum fimbriatum (1-9): 1. habit, X 1 ; 2. stem and fascicle, X 5; 3. stem in cross section, X 120;
4. branch in cross section, x 1 50; 5. stem leaf, x 24; 6. branch leaf, x 24; 7. branch leaf in cross section, x 250;
8. stem leaf areolation, x435; 9. branch leaf areolation, x435. S. obtusiusculum (10-18): 10. habit, xl; 11.
stem and fascicle, x 5; 12. stem in cross section, x 130; 13. stem leaf, x 20; 14. branch leaf, x 20; 15. stem leaf
in cross section (central part), X 150; 16. branch leaf in cross section (central part), X 150; 17. stem leaf aerola-
tion, x 435; 18. branch leaf areolation, X435. S. strictum subsp. pappeanum (19-27): 19. habit, xl; 20.
stem and fascicle, x5; 21. stem in cross section, x 120; 22. stem leaf, X20; 23. branch leaf, x20; 24. stem leaf
in cross section (central section), x 240; 25. branch leaf in cross section (at margin), x 240; 26. stem leaf areola-
tion, x 300; 27. branch leaf areolation, x435. (1-9, Wager 1525; 10-18, Wilman 527; 19-27, Esterhuysen
Sphagnaceae
25
26
Sphagnaceae
This widespread species is known from Europe,
Asia, North America, Australia, southern South
America and Southern Africa. Sphagnum fimbriatum
is rare in the Flora area, collected only once at Belfast
in the eastern Transvaal. Map 1.
Voucher: Wager 1057.
Sphagnum fimbriatum was originally reported for
Southern Africa (Dixon & Wager, 1930) from George
in the Cape. However, Wager’s own specimen
(PRE) and a specimen sent to the National Her-
barium by Wager both indicate that the collecting
locality is Belfast, Transvaal. If the Transvaal loca-
tion is correct it is possible that the species was
introduced from Europe during trout introduction
in the streams of that area.
2. Sphagnum obtusiusculum Lindb. ex
Warnst. in Hedwigia 29: 196 (1890); Warnst.,
Sphag. Univ. 90 (1911). Syntypes: Madagas-
car, Pollen & Van Dam s.n. ; Besson s.n. ;
Reunion, Richards 683; Rodriguez s.n.;
Chauvel s.n.; Mauritius, s. coll, in herb.
Renauld & Cardot.
Sphagnum laceratum sensu Garside in J1 S. Afr.
Bot. 15: 71 (1949), non C. Mull. & Warnst. (1897).
Plants medium to large, pink, becoming
yellowish with age, brownish below; terri-
colous. Stems to 60 mm high; internal cylinder
pink, hyalodermis efibrillose, pores scattered;
in section inner cells of internal cylinder large,
outer 3-4 rows smaller, strongly incrassate,
hyalodermis 2-3 cells thick. Fascicles of 3-4
monomorphic spreading branches; internal
cylinder pink, hyalodermis efibrillose, pores
1-2 per cell. Stem leaves oblong, 1 ,0-1 ,6 mm
long; apex obtuse to rounded, serrate;
margins erect, border of 4-6 rows of narrow
cells extending to mid-leaf; leucocysts efibril-
lose or occasionally very weakly fibrillose in
upper leaf, pores rare, some cells with single
pore; in section chlorocysts rectangular to
triangular with wider exposure on ventral
surface, leucocysts bulging dorsally. Branch
leaves ovate-acuminate, 1,2-1, 6 mm long;
apex acute, with 1-2 teeth; margins incurved
above, border of 2-3 rows of narrow cells
extending to near apex; leucocysts regularly
fibrillose, pores few, 1-3 per cell; in section
chlorocysts triangular to trapezoid with wider
exposure on ventral surface, leucocysts bulgi ng
dorsally.
Sporophyte not known. Fig. 3: 10-18.
Sphagnum obtusiusculum is found on Mada-
gascar, Reunion and Mauritius. In Southern Africa
the species is infrequently collected on damp soil or
rock in the southern and southwestern Cape. Map 2.
Map 2. — • Sphagnum capense
X Sphagnum obtusiusculum
Vouchers: Brenan M2783; Esterhuysen 26974;
Muir PRE-CH4138.
The most striking character of S. obtusiusculum
is its vivid pink colour. Several of the Southern
African species are tinged or partly coloured with red
or purple hues, but in S. obtusiusculum the exposed
parts of the plants are markedly coloured throughout;
the lower parts of the stem are brownish.
3. Sphagnum strictum Sull. subsp. pap-
peanum (C. Mull.) Eddy in Bull. Br. Mus. nat.
Hist. 5 (7): 433 (1977). Type: Cape, Swellen-
dam, Pappe s.n., 1838.
Sphagnum pappeanum C. Miill., Syn. Muse. 1 : 101
(1848); Warnst., Sphag. Univ. 151 (1911); Sim, Bryo.
S. Afr. 131 (1926); Garside in J1 S. Afr. Bot. 15: 71
(1949).
Plants large to robust, pale yellow to
yellow-green, occasionally reddish, brownish
below; terricolous. Stem 60-180 mm long;
internal cylinder green to yellow-brown,
hyalodermis efibrillose, pores ± 1 per cell; in
section cells of internal cylinder weakly
thickened, smaller toward margins, hyaloder-
mis 2-3 cells thick. Fascicles of 5(6) dimorphic
branches: 2-3 spreading, 2-3 pendent; inter-
nal cylinder green to yellowish, hyalodermis
efibrillose, ±1 pore per cell. Stem leaves
triangular, 0, 8-1,0 mm long; apex obtuse,
erose-denticulate; margins erect, border indis-
tinct, 6-8 cells wide below; leucocysts efibril-
lose or occasionally some cells weakly
fibrillose at apex, pores few, frequently with
resorption gaps; in section chlorocysts oval,
enclosed or just exposed on dorsal surface,
Sphagnaceae
27
leucocysts weakly convex on both surfaces.
Branch leaves ovate to oval, acuminate,
2, 5-3,0 mm long; apex truncate, dentate;
margins erect to incurved above, border of
single row of narrow cells extending to apex,
in section with resorption furrow; leucocysts
regularly fibrillose, pores scattered along
commissures; in section chlorocysts narrowly
oval, enclosed or just exposed dorsally, com-
missure wall papillose, leucocysts convex on
both surfaces.
Perichaetia on short lateral branch near
apex; leaves oval, 4 mm long, convolute, apex
rounded, serrate. Pseudopodia to 8 mm long;
capsules oval, 1,5 mm long, red-brown; oper-
culum and calyptra not seen; spores bulging
triangular, 35//m, coarsely granulate. Fig. 3:
19-27.
Sphagnum strictum subsp. pappeanum occurs in
Central America and the West Indies, eastern and
Southern Africa, the East African Islands, Celebes
and New Guinea. In Southern Africa, the subspecies
has been collected in the southern and southwestern
Cape, Natal and the eastern Transvaal. Map 3.
Map 3. — • Sphagnum pycnocladulum
x Sphagnum pappeanum
Vouchers: Cholnoky 843; Magill 6070; Van der
Schijff A52\ ; Taylor 1136b.
This taxon is identified by its multi-layered hyalo-
dermis, large oval-acuminate branch leaves and the
resorption furrow along the margins of the branch
leaves. The subspecies pappeanum differs from the
typical subspecies, which occurs in North America
and Europe, by its larger leaves, enclosed, oval
chlorocysts, and coarser papillation of the commis-
sure walls (cf. Eddy, 1977).
4. Sphagnum perichaetiale Hampe in
Linnaea 20; 66 (1847); Warnst., Sphag. Univ.
486 (191 1); Eddy in Bull. Br. Mus. nat. Hist.
5(7): 380 (1977). Type: Brazil (BM, holo.).
Sphagnum balfourianum Warnst. in Hedwigia 30:
153 (1891); Warnst., Sphag. Univ. 470 (1911);
Garside in J1 S. Afr. Bot. 15: 67 (1949), fide Eddy
(1977). Syntypes: Mauritius, Balfour s.n. (NY);
Cape, Rehmann s.n. (H-BR!).
Sphagnum marlothii Warnst. in Sphag. Univ. 471
(1911); Garside in J1 S. Afr. Bot. 15: 68 (1949), fide
Eddy (1977). Type: Cape, Table Mountain, Marloth
s.n.
Plants large to robust, pale green, occa-
sionally slightly pinkish, brownish below;
terricolous or saxicolous. Stems 80-140 mm
high; internal cylinder red-brown, hyaloder-
mis efibrillose to weakly fibrillose, pores ± 1
per cell; in section cells of internal cylinder
thickened, outer 3-4 rows much smaller,
strongly thickened, hyalodermis in 3 layers.
Fascicles of 3-4 dimorphic branches: 2-3
spreading, 1-2 pendent; internal cylinder red-
brown, hyalodermis fibrillose, pores scattered
or 1 per cell. Stem leaves variable in shape;
broadly elliptical to weakly obovate, 1 ,2-2,0
mm long; apex rounded, serrate or weakly
notched; margins unbordered; leucocysts
regularly fibrillose above base, occasionally
weakly so, pores few, irregularly placed along
commissures; in section chlorocysts oval to
trapezoid with wider exposure on ventral
surface, leucocysts mostly flattened ventrally,
strongly bulging dorsally. Branch leaves
broad, ovate to oval, 2,0-2, 8 mm long; apex
obtuse, cucullate; margins entire, erect, un-
bordered; leucocysts strongly fibrillose
throughout, pores few, scattered along com-
missures; in section chlorocysts triangular to
trapezoid or oval with wider exposure on
ventral surface, leucocysts flattened ventrally,
strongly bulging dorsally.
Sporophytes not known from Southern
Africa. Fig. 4: 1-12.
This widespread species is known from North
and Central America, northern South America,
Africa south of the equator. East African Islands,
southern Asia, and Australia. In Southern Africa
S. perichaetiale is found in the southwestern and
southern Cape and southern Natal. Map 4.
Voucher: Esterhuysen 15839.
In addition to the robust plants and large ovate
branch leaves, the presence of fibrils in the hyaloder-
mis of branches and occasionally stems, will help to
place specimens of S. perichaetiale.
28
Sphagnaceae
Fig. 4. — Sphagnum perichaetiale: 1. habit, xl;
2. stem and fascicle, x5; 3. stem in surface view,
x25; 4. stem in cross section, x80; 5. branch in
surface view, x 50; 6. branch in cross section, x 170;
7. stem leaf, x 15; 8. branch leaf, x 15; 9. stem leaf in
cross section (central part), x220; 10. branch leaf
cross section (central part), x220; 11. stem leaf
leucocyst, x435; 12. branch leaf leucocyst, X435.
(1-12, Esterhuysen 15839).
5. Sphagnum africanum Welw. & Dub. in
Mem. Soc. Phys. Hist. nat. Geneve 21: 216
(1870); Warnst., Sphag. Univ. 421 (1911).
Type; Angola, Huilla, Welwitsch 77 (G!).
Sphagnum oligodon C. Mull, in Flora, Jena 70: 412
(1887); Warnst., Sphag. Univ. 363 (1911). Type:
Natal, Inanda, Rehmann 14(PRE1).
Sphagnum rehmannii Warnst. in Hedwigia 30: 16
(1891); Warnst., Sphag. Univ. 372 (1911); Sim,
Bryo. S. Afr. 132 (1926). Syntypes: Natal, Rehmann
s.n.; Gueinzius s.n. (NY); Transvaal, MacLea s.n.
(Rehmann 431, PRE!).
Sphagnum transvaaliense Warnst. in Hedwigia 30:
32 (1891). Type: Transvaal, Spitzkop to Lydenburg,
Wilms s.n. (Gl).
Sphagnum oligodon var. bachmannii Warnst.,
Sphag. Univ. 363 (1911). Type: Transkei, Pondoland,
Bachmann 5 (H-BR).
Sphagnum oligodon var. beyrichii Warnst., Sphag.
Univ. 364 (1911). Type: Transkei, Pondoland,
Beyrich 25 (H-BR).
Plants medium to large (robust), pale
green to yellow-green, brownish below ; saxi-
colous or terricolous. Stems 35-120 mm high;
internal cylinder green, hyalodermis efibril-
lose, pores few; in section cells of internal
cylinder not thickened or only weakly so,
hyalodermis 1 cell thick. Fascicles of 3-4
dimorphic branches: 2-3 spreading, 1-2
pendent; internal cylinder greenish, hyaloder-
mis efibrillose, pores weakly defined. Stem
leaves broad, ovate to oblong, 2, 4-2, 7 mm
long; apex obtuse, serrulate; margins erect,
border of 2-4 rows of narrow cells extending
to near apex; leucocysts regularly fibrillose,
pores numerous, regularly placed along com-
missure, in section chlorocysts rectangular to
oval, equally exposed on both surfaces,
leucocysts weakly bulging on both surfaces.
Branch leaves broad, oval or occasionally
ovate, 1,8-2, 8 mm long; apex rounded,
serrate; margins incurved above, border of
1-2 rows of narrow cells to near apex; leuco-
cysts regularly fibrillose, pores numerous
along commissures; in section chlorocysts
rectangular to oval, exposure equal, cells very
thin-walled, leucocysts weakly bulging on
both surfaces.
Sporophyte not known. Fig. 5: 19-27.
The species was described from Angola and has
recently been discovered in Southern Africa. In the
Flora area the species is found in streams, pools or
very wet areas of the western, central and eastern
Transvaal, Swaziland, Zululand, Natal, Transkei
and southern Cape. Map 4.
Vouchers: Arnold 1238; Cholnoky 259, 876;
Hilliard & Burtt 11793; Magill 5271; Rankin 120;
Smook 859; Van Rooy 170; Venter 2999; Von Breiten-
bach 54.
Sphagnum africanum is identified by its single-
layered hyalodermis, large stem leaves, broadly oval
branch leaves with rounded, serrate apices and
chlorocyst in section being rectangular and equally
exposed on both surfaces. The species is very closely
Sphagnaceae
29
?
rr
FFttH
ttnua-
tLJ [I
—
—
-
H+m
+FFS
a
2
-
-
"nIjTTT
-
r
..
*
Map 4. — • Sphagnum africanum
x Sphagnum perichaetiale
related to S. truncatum and the two species occasion-
ally prove difficult to separate. Differences in branch
leaf shape, development of branch leaf apices, chloro-
cyst shape and exposure in section, and to a lesser
degree distribution will generally separate specimens.
6. Sphagnum truncatum Hornsch. in Lin-
naea 15: 114 (1841); Warnst., Sphag. Univ.
386 (1911); Sim, Bryo. S. Afr. 132 (1926);
Garside in J1 S. Afr. Bot. 15: 72 (1949). Type:
Cape, Du Toit’s Kloof Mountains, Drege s.n.
(G!).
Sphagnum coronatum C. Miill. in Flora, Jena 70:
412 (1887); Warnst., Sphag. Univ. 306 (1911); Sim,
Bryo. S. Afr. 132 (1926). Type: Cape, Montagu Pass,
Rehmann 9 (PRE!).
ISphagnum fluctuans C. Miill. in Flora, Jena 70:
414 (1887). Type: Cape, Gnadenthal, Breutel s.n.;
vide Warnst., Sphag. Univ. 309 (1911).
Sphagnum convolutum Warnst. in Hedwigia 29:
217 (1890). Sphagnum marginatum var. convolutum
Warnst., Sphag. Univ. 309 (1911). Syntypes: Cape,
Table Mountain, MacOwan s.n., 1886; Simonstown,
Wright s.n. (FH).
Sphagnum marginatum Warnst. in Hedwigia 30: 28
(1891). Type: Cape, Sonderend, Breutels.n. (BM; PC).
Sphagnum coronatum var. cuspidatum Warnst.,
Sphag. Univ. 306 (1911). Type: Cape, Worcester,
Rehmann 10, p.p.
Sphagnum marginatum var. diversifolium Warnst.,
Sphag. Univ. 310 (1911). Type: Cape, Sonderend,
Breutel s.n. (PC).
Sphagnum oxycladum Warnst. in Hedwigia 30: 15
(1891); Warnst., Sphag. Univ. 306 (1911). Type:
Cape, Worcester, Rehmann 10, p.p.
Plants medium to large, pale green to
yellow-green, occasionally reddish, brown
below; terricolous or saxicolous. Stems
35-110 mm high; internal cylinder brownish
to green, hyalodermis efibrillose, pores weakly
defined, ± 1 per cell; in section inner cells
large, thin-walled, outer cells smaller, fre-
quently incrassate, coloured, occasionally
only weakly thickened, hyalodermis 1 cell
thick. Fascicles of 4-5 dimorphic branches:
2-3 spreading, 1-2 pendent; internal cylinder
brownish, hyalodermis efibrillose, pores
weakly defined. Stem leaves oblong, 1, 8-2,0
mm long; apex obtuse; margins plane, border
of 2-4 rows of narrow cells extending to near
apex; leucocysts regularly fibrillose, pores
few, scattered along commissures; in section
chlorocysts trapezoid with wider exposure on
dorsal surface, rarely elliptical, equally ex-
posed, leucocysts bulging ventrally or occa-
sionally weakly bulging on both surfaces.
Branch leaves broadly lanceolate to ovate-
acuminate, 2, 0-4, 2 mm long; apex truncate,
toothed; margins erect, border of 1-2 rows of
narrow cells extending to upper leaf; leuco-
cysts regularly fibrillose, pores few, scattered
along commissures; in section chlorocysts
trapezoid with wider exposure on dorsal
surface, leucocysts bulging ventrally.
Sporophyte not known. Fig. 5: 1-9.
Endemic to Southern Africa, S. truncatum is
most common in the southwestern and southern Cape.
A few specimens have also been collected in pools in
the Transkei and in central Transvaal. Map 1.
Vouchers: Cholnoky 302, 1077; Emdon 4;
Schelpe 7838; Stokoe 9490.
The single-layered hyalodermis, large lanceolate
branch leaves and chlorocysts trapezoid in section,
will identify S. truncation. This species is similar to S.
africammv, for differences see note under that species.
Variation in S. truncatum is expressed in the size
and shape of branch leaves, shape of chlorocysts in
section and degree of thickening of the outer cell in
the internal cylinder of the stem.
7. Sphagnum pycnocladulum C. Miill. in
Flora, Jena 70: 420 (1887); Warnst., Sphag.
Univ. 174 (1911); Sim, Bryo. S. Afr. 131
(1926). Type: Cape, Montagu Pass, Rehmann
s.n., 1875.
Sphagnum pycnocladulum var. fuscescens Warnst.,
Sphag. Univ. 176 (1911). Type: Cape, Montagu
Pass, Rehmann 13 (NH; PRE!).
Sphagnum pycnocladulum var. viride Warnst.,
Sphag. Univ. 176 (1911). Type: Cape, Montagu
Pass, Rehmann 17 (PRE!).
30
Sphagnaceae
Sphagnaceae
31
Plants medium to large, yellow-green to
yellowish, occasionally reddish, brown below;
terricolous or saxicolous. Stems 50-120 mm
high; internal cylinder brownish, hyalodermis
efibrillose, pores weakly defined, ± 1 per cell ;
in section inner cells large, thin-walled,
becoming smaller and incrassate toward
margin of internal cylinder, hyalodermis 2-3
cells thick. Fascicles of 5-6 monomorphic
branches; internal cylinder brownish, hyalo-
dermis efibrillose, pores scattered to 1 per cell.
Stem leaves broad, oblong to spathulate,
0,9- 1,2 mm long; apex obtuse; margins
plane, bordered by 2-4 rows of narrow cells
to near apex; leucocysts essentially efibrillose,
pores large, numerous, confined to commis-
sures, absent in leaf base; in section chloro-
cysts oval, totally enclosed or occasionally
equally exposed on both surfaces; leucocysts
slightly bulging on both surfaces. Branch
leaves ovate, 0,6-0, 8 mm long; apex acute;
margins involute above, border of narrow
cells 1-2 cells wide, extending to near apex;
leucocysts fibrillose, although frequently
weakly so, occasionally efibrillose, pores
numerous along commissures; in section
chlorocysts oval, totally enclosed or some-
times equally exposed on both surfaces,
leucocysts slightly bulging dorsally.
Perichaetia terminal ; leaves broadly oval,
2,7 mm long; apex broadly acute, somewhat
irregular; leucocysts fibrillose to efibrillose.
Pseudopodia 4, 5-5,0 mm long; capsule
exserted, dark red-brown, mouth wide-
spreading when deoperculate; operculum and
calyptra not seen; spores bulging triangular,
35 pm, tetrad scar obvious, finely granulose.
Fig. 5: 10-18.
In Southern Africa the species is presently only
known from the Cape Province but it has also been
reported from Ruwenzori and Usambara in eastern
Africa. Map 3.
Vouchers: Cholnoky 305; Esterhuysen 27350;
Garside 6293 ; Magill 6297.
Sphagnum pycnocladulum is in many ways very
similar to S. capense. Differences are most obvious
in the number of branches in the fascicles, leaf size
and, to a lesser degree, shape and the development of
fibrils in leaf leucocysts. The stem leaf fibrils in S.
capense are regular and well developed, while in S.
pycnocladulum the stem leaf leucocysts are practically
efibrillose. In addition the branch leaf fibrils of the
latter are generally not as regular or as well developed.
The two species also have different distribution
areas. Southern African collections of S. pycnocladu-
lum are presently known only from the southern and
southwestern Cape. Sphagnum capense is found in
practically the same areas and is also known from
Transkei, Natal and eastern Transvaal. Outside
Southern Africa, S. pycnocladulum occurs throughout
eastern Africa, while S. capense is found on the East
African Islands. Considering the variability expressed
by both species in Southern Africa, S. pycnocladulum
is only provisionally maintained at the species level.
8. Sphagnum capense Hornsch. in Linnaea
15: 113 (1841); Warnst., Sphag. Univ. 427
(191 1); Sim, Bryo. S. Afr. 130 (1926); Garside,
p.p. in J1 S. Afr. Bot. 15: 75 (1949); Eddy in
Syst. Ass., sp. vol. 14: 109-121 (1979). Syn-
types: Cape, Table Mountain, Ecklon s.n. ;
Devil’s Kloof, Drege s.n.
Sphagnum austromolle C. Miill. in Flora, Jena 70:
419 (1887). Sphagnum capense var. austromolle (C.
Miill.) Warnst., Sphag. Univ. 430 (1911). Syntypes:
Cape, Table Mountain, Rehmann s.n., Nov. 1875;
Devil's Peak, Rehmann s.n., Oct. 1876; Montagu Pass,
Rehmann s.n.
Sphagnum mollissimum C. Miill. in Flora, Jena 70:
418 (1887). Sphagnum capense var. mollissimum (C.
Miill.) Warnst., Sphag. Univ. 430 (1911). Syntypes:
Cape, Table Mountain, Rehmann s.n., 1875; Spielhaus
s.n., 1877; Montagu Pass, Rehmann s.n.; Stinkwater,
Rehmann s.n.
Sphagnum panduraefolium C. Miill. in Flora, Jena
70: 418 (1887); Warnst., Sphag. Univ. 299 (1911);
Sim, Bryo. S. Afr. 131 (1926); Garside in J1 S. Afr.
Bot. 15: 74(1949). Syntypes: Cape, Table Mountain,
Rehmann s.n., 1875; Stinkwater, Rehmann 16 (BOL!;
PRE!).
Fig. 5. — Sphagnum truncatum (1-9): 1. habit, x 1 ; 2. stem and fascicle, x 5; 3. stem in cross section, x 100;
4. stem leaf, x20; 5. branch leaf, x20; 6. stem leaf in cross section (central part), x220; 7. branch leaf cross
section (central part), x220; 8. stem leaf areolation, x435; 9. branch leaf areolation, x435. S. pycnocladulum
(10-18): 10. habit, xl; 11. stem and fascicle, x5; 12. stem in cross section, x 1 25 ; 13. stem leaf, x20; 14.
branch leaf, x 20; 15. stem leaf in cross section (central part), x 220; 16. branch leaf in cross section (at margin),
x220; 17. stem leaf areolation, x435; 18. branch leaf areolation, x435. S. africanum (19-27): 19. habit, xl;
20. stem and fascicle, x 5; 21. stem in cross section, x 100; 22. stem leaf, x 20; 23. branch leaf, x 20; 24. stem
leaf in cross section (central part), x220; 25. branch leaf in cross section (central part), x220; 26. stem leaf
areolation, x435; 27. branch leaf areolation, x435. S. capense (28-36): 28. habit, xl; 29. stem and fascicle,
x5; 30. stem in cross section, x 190; 31. stem leaf, x20; 32. branch leaf, x20; 33. stem leaf in cross section
(at margin), x220; 34. branch leaf in cross section (at margin), x220; 35. stem leaf areolation, X435; 36.
branch leaf areolation, x435. (1-9, Levyns 24066; 10-18, Esterhuysen 15427; 19-27, Vorster 509; 28-36, Ester-
huysen 15431).
32
Sphagnaceae
Sphagnum capense var. multiporosum Warnst.,
Sphag. Univ. 428 (1911). Syntypes: Cape, Cape Town,
Ecklon s.n., 1827; Ecklon & Zeyher s.n., 1863;
Rehmann 16c, 433 (PRE!), 434b (PRE!); Wilms 2629;
Laux s.n.; Marloth s.n., 1902; Devil’s Peak, Rehmann
s.n.
Sphagnum beyrichianum Warnst., Sphag. Univ.
385 (1911). Type: Transkei, Pondoland, Beyrich s.n.
(H-BR).
Plants medium to large, yellow-green to
light green, occasionally reddish, brownish
below; terricolous or saxicolous. Stems
40-110(-220) mm high; internal cylinder
brownish, hyalodermis efibrillose, pores
weakly defined, ±1 per cell; in section inner
cells large, thin-walled, becoming smaller,
incrassate toward margin of internal cylinder,
hyalodermis 2-3 cells thick. Fascicles of 3-4
dimorphic branches: 2-3 spreading, 1 pen-
dent; internal cylinder brownish, hyalodermis
efibrillose, pores ±1 per cell. Stem leaves
broad, oval to oblong, rarely obovate,
1,0-1, 7 mm long; apex rounded to acute;
margins plane to erect above, border of 1-2
rows of narrow cells extending to near apex;
leucocysts regularly fibrillose, pores numerous
along commissures, occasionally with free
pores; in section chlorocysts oval, enclosed,
leucocysts weakly bulging on both surfaces.
Branch leaves ovate, 0,8- 1,2 mm long; apex
acute; margins incurved above, border of 1-2
rows of narrow cells extending to apex;
leucocysts regularly fibrillose, pores numerous
on commissures, occasionally pores few, fre-
quently with free pores; in section chlorocysts
oval, enclosed, leucocysts weakly bulging on
both surfaces.
Sporophytes not seen. Fig. 5: 28-36.
Sphagnum capense is infrequently collected on
wet soil or rock in mountains of the southern and
southwestern Cape, Transkei, Natal and eastern and
central Transvaal. The species has also been collected
in Madagascar and Reunion. Map 2.
Vouchers: Cholnoky 1100; Esterhuysen 15431;
Kluge 1053; Magi II 6316; Vorster 461a.
This relatively small-leaved species is clearly
distinct from all other Southern African species
except S. pycnocladulum; see note on differences
under that species.
Variation in the Southern African specimens of
S. capense is most obvious in leucocyst ornamenta-
tion. The pores of the leucocysts are generally
numerous, occurring all along the commissure and
are also frequently free on the surface; however,
specimens have been seen with only a few, scattered
pores per cell. The fibrils of the leucocysts are typi-
cally regular and well developed. Careful examination
of leaves of a few specimens, however, reveal some
cells with weak or irregular fibrils among the more
typical cells.
33
ANDREAEACEAE
Plants small to large, forming patches or cushions, red-brown to green-black; saxicolous,
rarely semi-aquatic. Stems slender, generally erect; in section central strand absent, cortical
cells weakly incrassate. Leaves appressed dry, spreading to squarrose wet; ovate to elliptical,
occasionally constricted above base; apex obtuse to subulate. Costa present or absent; in
section cells not strongly differentiated. Laminal cells small, quadrate to rectangular, incrassate,
dorsal surface occasionally papillose.
Monoicous or dioicous. Perigonia on short lateral branches, gemmate. Perichaetia termi-
nal; leaves larger, convolute. Seta absent, sporophyte elevated at maturity by pseudopodia;
capsule elliptical to ovate-oblong, lacking peristome and operculum, dehiscing through 4-8
longitudinal valves; spores round, yellowish.
The family contains a single genus, Andreaea. with c. 100 species.
ANDREAEA
Andreaea Hedw., Spec. Muse. 47 (1801); Broth, in Natiirl. PfIFam. 10: 129 (1924); Sim,
Bryo. S. Afr. 133 (1926); Grout, Moss. FI. N. Amer. 1: 1 (1936); Sainsb., N. Zeal. Moss. 18
(1955); Schultze-Motel in Willdenowia 6: 25 (1970); Smith, Moss FI. Brit. Irel. 79 (1978).
Type species: A. rupestris Hedw.; vide Britt. & Emer. in N. Amer. FI. 15: 35 (1913).
With characters of the family.
Andreaea forms blackish or reddish patches on bare rock at high altitudes throughout the world. In
Southern Africa, Andreaea is found on granite near the summits of the higher mountains of the Cape, Lesotho,
Natal and the Orange Free State. One exception is the presence of two of the species on Table Mountain.
When fruiting the genus is unmistakable. The capsules are without operculum or peristome; dehiscence
occurs through the splitting of the capsule along 4-8 longitudinal valves.
1 Leaves costate:
2 Plants large; leaves broadly oval to ovate; costa broad below, not reaching upper leaf 2. A. nitida
2 Plants small; leaves narrowly ovate-subulate; costa filling subula l.A. subulata
1 Leaves ecostate:
3 Leaves unistratose 3. A. rupestris
3 Leaves bistratose 4. A. bistratosa
1. Andreaea subulata Harv. ex Hook, in
Hooker’s Icon. PI. 3: 201 (1840); Broth, in
Natur!. PflFam. 10: 130 (1924); Sim, Bryo. S.
Afr. 134 (1926); Schultze-Motel in Willde-
nowia 6: 77 (1970); De Sloover in Bull. Jard.
bot. nat. Belg. 47: 156 (1977). Type: Cape,
Table Mountain, The Port, Harvey s.n., 21
Mar. 1837 (BM, holo.!).
Plants medium-sized, loosely caespitose,
brown-green to black-green or red-brown;
saxicolous. Stems 10-20 mm long, irregularly
branched; in section round, central strand
absent, inner cortical cells in 3 rows, slightly
thickened, outer cortical cells in 3 rows,
smaller, incrassate. Leaves falcate-secund to
almost circinate wet or dry; oval to oblong,
long-subulate, 2 , 0-2 , 5(-3 , 0) mm long; mar-
gins plane, entire. Costa wide, filling subula;
in proximal section flattened, cells in 2 rows,
undifferentiated, strongly thickened; lamina
unistratose; in distal section reniform, inter-
nal cells in 2-4, rows, small strongly thicken-
ed, dorsal and ventral surface cells thin-
walled; lamina bistratose. Upper laminal cells
rounded-quadrate, incrassate, bulging dor-
sally, smooth ventrally; basal juxtacostal
cells rectangular.
Perichaetia terminal; leaves sheathing,
convolute, oval, short-acuminate to acute,
3, 0-3, 5 mm long. Pseudopodia light green, 2
mm long; capsules immersed to emergent,
elliptical-apiculate, 1,0-1, 2 mm long, red-
34
Andreaeaceae
Fig. 6.— Andreaea subulata (1-12): 1. habit,
xl; 2. habit, x5; 3. stem cross section, xl70; 4.
leaves, x40; 5. leaf in proximal cross section, xl70;
6. leaf in distal cross section, xl70; 7. leaf base,
xl70; 8. leaf areolation, x640; 9. leaf apex, xl70;
10-11. perichaetial leaves, x40; 12. capsule, dehis-
cent and dry, xlO; A. nitida (13-21): 13. habit,
xl; 14. part of stem, x5; 15. stem in cross section,
x 130; 16. leaf, x40; 17. leaf in cross section, x220;
18. cells at leaf base, x 170; 19. leaf areolation, x640;
20. leaf apex, xl30; 21. spore xl90. (1—12, Sim
9223; 13-21, Esterhuysen 18475).
Andreaeaceae
35
brown, dehiscence slits extending from base to
apex; spores round, 25-35 p m, yellow,
reticulate-papillose. Fig. 6: 1-12.
Originally described from a specimen collected
by Harvey on Table Mountain, A. subulata has
subsequently been collected on the East African
Islands and in Australia, New Zealand and southern
South America. The species has been re-collected
several times in the Flora area, but only on Table
Mountain. Map 5.
Vouchers: Esterhuysen 18599 (BOL); Sim 9223;
Wager 104.
The small size of the plants and the costate,
ovate-subulate leaves will place A. subulata. Occa-
sionally high-altitude, rock-growing specimens of
Grimmia apocarpa have been mistaken for this
species. These specimens are also frequently small
and reddish brown, but the larger leaves with broader
apices and sinuolate leaf cells should separate these
specimens.
2. Andreaea nitida Hook. f. & Wils. in J.
Bot., Lond. 3: 535 (1844); Broth, in Natiirl.
PflFam. 10: 131 (1924); Schultze-Motel in
Willdenowia 6 : 89 (1970) ; Schelpe in J1 S. Afr.
Bot. 41: 37 (1975). Type: Lord Auckland’s
Islands, Hooker s.n. (BM, holo. !).
Plants large, loosely tufted, reddish
brown to black-green; saxicolous, semi-
aquatic. Stems 20-120 mm long, branching;
in section round, central strand absent, inner
cortical cells in 5-6 rows, with thickened
corners, outer cortical cells in 3 rows, smaller,
strongly thickened. Leaves somewhat fragile,
extremely variable in size, lower leaves small,
becoming larger above, erect-spreading wet or
dry; broadly elliptical, ( 1 — )2— 3 mm long, to
1 ,5 mm wide; apex broadly obtuse, abruptly
apiculate; margins plane, entire. Costa broad
below, tapering to mid-leaf, laterally spurred,
cells in surface view elongate; in section cells
weakly differentiated, slightly smaller, thick-
walled. Upper laminal cells variable, quadrate,
rectangular or triangular, weakly thickened,
smooth.
Perichaetia terminal; leaves weakly diffe-
rentiated from upper leaves. Pseudopodia 3-5
mm long, blackish; capsule exserted, elliptical,
2-3 mm long, dark red-black, dehiscence slits
extending from mid-capsule to apex; spores
round, 20-30 //m; brownish yellow, weakly
papillose. Fig. 6: 13-21.
Rare in Southern Africa, A. nitida is more com-
mon in Australia, New Zealand, South America and a
few Antarctic Islands. The species is presently known
only from the Hexberg in the Cold Bokkeveld Moun-
tains of the southwestern Cape. Map 5.
Map 5. — • Andreaea bistratosa
x Andreaea subulata
A Andreaea nitida
Voucher: Esterhuysen 18475 (BOL).
Andreaea nitida is identified by its broadly
elliptical leaves and wide, spurred costa which tapers
to mid-leaf. The size of the leaves increases consider-
ably up the stem, especially on fertile plants. The
size of the plants and leaf shape are very distinct
from the other Southern African species of Andreaea.
and the species is unlikely to be confused with other
taxa after careful examination.
3. Andreaea rupestris Hedw., Spec. Muse.
47 (1801); Scott & Stone, Moss. S. Aust. 62
(1976); Smith, Moss FI. Brit. Irel. 81 (1978).
Type: Europe.
Andreaea petrophila [Ehrh., 1784] Fuernr. in Flora,
Jena 10: 30 (1827); Broth, in Natiirl. PflFam.
10: 129 (1924); Sim, Bryo. S. Afr. 133 (1926). Type:
Europe.
Plants small, forming cushions, red-
brown to black-green; saxicolous. Stems 5-20
mm high, irregularly branched; in section
round, central strand absent, inner cortical
cells in 2-3 rows, corners thickened, outer
cells in 2 rows, smaller, round, incrassate.
Leaves appressed or with spreading apices
dry, erect-spreading wet; ovate to oblong-
lanceolate, occasionally constricted above
base, 0,3-1, 5 mm long; apex weakly cucul-
late, acute to obtuse; margins entire, in-
curved; lamina in section unistratose. Costa
absent. Upper laminal cells quadrate, incras-
sate, frequently with strongly thickened
corners, smooth to strongly papillose on
dorsal surface; basal cells linear to rectangu-
lar, irregularly thickened; basal marginal
cells short-rectangular to quadrate.
36
Andreaeaceae
Autoicous. Perichaetia terminal; leaves
elliptical, 2 mm long; apex obtuse. Pseudo-
podia 1-2 mm long, pale green; capsule just
exserted, elliptical, to 1 mm long, dark red-
brown; dehiscence slits extending from mid-
capsule to near apex; spores round, 20-25
pm, yellow, weakly papillose. Fig. 7: 11-24.
Very widespread in distribution, A. rupestris
is also the most commonly collected Andreaea in
Southern Africa. Growing on bare rock, at high
altitude, this species is known from mountains of the
southwestern, central and eastern Cape and the
Drakensberg of Lesotho, Natal and the Orange
Free State. Map 6.
Vouchers: Brenan M2774; Ellis 7; Esterhuysen
34592; Hilliard & Burtt 10497; Magill 4416; Schelpe
2114; Smook 1112.
In Southern Africa, A. rupestris is identified by
its generally red-brown colour, ecostate leaves and
frequently papillose leaf cells. The plants tend to
vary in size and leaf shape, but specimens are rarely
difficult to identify. A broader interpretation of A.
rupestris would undoubtedly encompass many of the
ecostate species of Andreaea on the sub-Antarctic
Islands.
Fig. 7. — Andreaea bistratosa (1-10): 1. habit, xl; 2. part of stem, x5; 3. stem in cross section, x300;
4. leaves, x30; 5. cells at leaf base, x 170; 6. leaf apex, x 170; 7. leaf in proximal cross section, x 170; 8. leaf
in distal cross section, X 170; 9. basal leaf cells, X640; 10. upper laminal cells at margin, x640. A. rupestris
(11-24): 11. habit, xl; 12. habit, x5; 13. stem in cross section, x300; 14-16. leaves, x40; 17. cells at leaf
base, x 170; 18. leaf apex, x 170; 19. upper laminal cells, x640 ; 20. basal leaf cells, x640; 21. leaf in proxi-
mal cross section, x 170; 22. leaf in distal cross section, x 170; 23. capsule, intact, x 10; 24. capsule, dehiscent
and dry, x 10. (1-10, Esterhuysen 20601 ; 11-24, Wager PRF.-CH12173).
Andreaeaceae
37
4. Andreaea bistratosa Magill, sp. nov.,
formis A. rupestris Hedw. similis sed foliis
anguste lanceolatis bistratosis et cellulis lami-
nae regularibus, infirme incrassatis differt;
speciebus austro-africanis aliis foliis ecostatis
differt.
Type: Cape, Schurweberg, on sandy wet
rock surface in cool gully, 1 400 m, Ester-
huysen 20601 (BOL, holo. ; MO; PRE).
Plants small, forming loose cushions,
blackish to brownish green; saxicolous. Stems
5-20 mm tall, irregularly branched; in section
round, central strand absent, inner cortical
cells in 3 rows, corners thickened, outer corti-
cal cells in 2 rows, smaller, incrassate. Leaves
appressed dry, widespreading above base wet,
crowded above, fragile on lower stem, gene-
rally only leaf bases remaining; lanceolate,
occasionally constricted above base, 1 ,0-1 ,5
mm long; apex acute; margins plane, entire;
lamina in section bistratose, cells becoming
slightly larger toward margins. Costa absent.
Upper laminal cells hexagonal to quad-
rate, weakly papillose dorsally, becoming
rhomboidal to rectangular below, weakly
thickened; basal cells long-rectangular, thin-
walled.
Dioicous. Perigonial buds terminal,
rounded; leaves broadly ovate-acute, convo-
lute; antheridia few. Perichaetia terminal;
leaves similar to vegetative leaves, papillose
dorsally; archegonia few. Sporophyte not
known. Fig. 7: 1-10.
Endemic to Southern Africa, A. bistratosa has
been collected on rock, above 1 400 m, in the moun-
tains of the southwestern Cape. Map 5.
Voucher: Esterhuysen 25507.
Similar to forms of A. rupestris, this species is
separated through its narrow, bistratose leaves and
its very regular, weakly thickened laminal cells.
Andreaea bistratosa is distinguished from the other
Southern African species by its ecostate leaves.
39
FISSIDENTACEAE
Plants minute to large, generally forming dense mats; terricolous, saxicolous, corticolous
or rarely aquatic. Stems erect or prostrate, generally simple; central strand present or absent.
Leaves distichous and complanate, larger above, equitant, generally long, narrow, occasionally
asymmetrical; Ungulate to lanceolate or linear; in proximal transverse section Y-shaped,
ventrally with 2 vaginant laminae clasping stem, dorsally with single lamina; apices acute to
rounded; margins elimbate or completely to partly limbate, smooth, crenulate or serrate.
Costae generally well developed, short-excurrent or ending below apex, rarely absent. Vaginant
laminae equal or unequal, extending | to | of leaf length; dorsal lamina tapering proximally to
insertion, frequently not reaching insertion or rarely decurrent. Lamina l cells mostly small,
quadrate to hexagonal, occasionally oblong-hexagonal or rhombic, flat to mammillose,
smooth or papillose.
Perichaetia terminal or lateral. Seta elongate, erect or flexuose; capsules erect to inclined,
urns short-cylindrical, symmetric or curved; peristome single, teeth 16, generally deeply cleft,
reddish to red-yellow; operculum rostrate; calyptra cucullate; spores small, generally round.
A family of about 830 species in a single genus, Fissidens. Three other segregate genera have been recog-
nized by some authors.
FISSIDENS
Fissidens Hedw., Spec. Muse. 152 (1801); Broth, in Natiirl. PflFam. 10: 144 (1924); Sim, Bryo. S.
Afr. 185 (1926) ; Grout, Moss FI. N. Amer. 1 : 7 (1936); Gangulee, Moss. E. India 447 (1971).
Lectotype species: F. bryoides Hedw., fide Britt, in Britt. FI. Bermuda 435 (1916).
Octodiceras Brid., Muscol. Recent. Suppl. 1 : 162 (1806); Smith, Moss FI. Brit. Irel. 204 (1978). Type species:
O . fissidentoides Brid.
Skitophyllum B. Pyl. in J. de Bot., ser. 2, 4: 133 (1814), nom. illeg.
Conomitrium Mont, in Annls Sci. nat. Bot., ser. 2, 8: 245 (1837), nom. illeg.
Moenkemeyera C. Miill. in Flora, Jena 69: 506 (1886); Broth, in Natiirl. PflFam. 10: 154 (1924). Type species:
M. mirabilis C. Miill.
With characters of the family.
The genus Fissidens comprises c. 830 currently recognized species. The genus has a worldwide distribution,
with major centres of described species in South America (243) and Africa (268). Seventy-three species have
been described or reported from the Flora area; of these Sim (1926) recognized 40 species. The number is here
reduced to 28.
1 Plants small to minute; stems without central strand; leaves lacking costa (Subgenus Aneuron ):
2 Leaves limbate; marginal leaf cells strongly differentiated, linear 1 . F. enervis
2 Leaves elimbate; marginal leaf cells quadrate to short-rectangular 2. F. usambaricus
1 Plants small to large; stems with central strand, sometimes weak; leaves with costa short or long (Sub-
genus Fissidens):
3 Leaf cells lax, elongate-hexagonal to fusiform; costa short, ending well below apex (Section Areofissi-
dens) :
4 Leaves 2-4 mm long; leaf cells 60-110 pm. long 3. F. wageri
4 Leaves 0 , 5-2 , 0 mm long ; leaf cells 20-50 pm long :
5 Leaves limbate; marginal cells elongate, leaf cells 20-32 pm long 5. F. capriviensis
5 Leaves elimbate; marginal cells somewhat smaller, leaf cells 30-50 pm long 4. F. splachnifolius
40
Fissidentaceae
3 Leaf cells smaller with ± thickened walls; costa extending to near apex:
6 Leaves completely or in part limbate:
7 Limbidia present on all laminae, strong or weak (Section Fissidens ):
8 Upper laminal cells regularly isodiametric, hexagonal or quadrate:
9 Vaginant laminae equal:
10 Laminal cells 4-6 pm long; limbidia 4-6 cells wide on proximal vaginant laminae
7. F. hoeegii
10 Laminal cells 8-12 pm long; limbidia generally not broader on proximal vaginant laminae
6. F. bryoides
9 Vaginant laminae unequal:
11 One blade of vaginant laminae terminating on lamina; limbidia intermarginal by 1-8 cells
in proximal vaginant laminae; laminal cells bulging, 4-6 pm long 8. F. rufescens
11 One blade of vaginant laminae terminating on costa; limbidia marginal throughout:
12 Leaves lanceolate; apex short-acuminate; laminal cells flat, quadrate to rhomboid,
15-18 long 10. F. simii
12 Leaves ligulate to elliptical; apex acute to broadly acute, cuspidate; laminal cells bulging,
quadrate, hexagonal or angular, 10-12 pm long 9. F. marginatus
8 Upper laminal cells generally longer than broad or heterogeneous, rectangular and angular:
13 Limbidia weakly differentiated on vegetative leaves, frequently only a few elongated cells pre-
sent on each lamina 12. F. stellenboschianus
13 Limbidia strongly differentiated:
14 Leaves 1 ,0-2,0 mm long; laminal cells lax, 35-42 /mi long 13. F. aciphyllus
14 Leaves 0,7-0, 8 mm long; laminal cells small, 10-17 pm long 11. F. curvatus
7 Limbidia present on vaginant laminae of perichaetial leaves, frequently also on vaginant laminae
of vegetative leaves, although usually weak to absent (Section Semilimbidium) :
15 Leaf cells smooth:
16 Leaves very short, 0, 3-0, 5 mm long, broadly ovate to ligulate; one blade of vaginant laminae
terminating on costa ; apex of leaves on sterile plants bent abruptly backwards
18. F. pygmaeus
16 Leaves longer, 0,6-1 ,2 mm long; one blade of vaginant laminae terminating on lamina:
17 Leaves lanceolate to elliptical, 0,6-0, 8 mm long; apex acute; limbidia on vaginant laminae
of perichaetial and subperichaetial leaves only; laminal cells irregular, short- to long-
hexagonal, 12-17 pm long 17. F. parvilimbatus
17 Leaves oblong, elliptical or asymmetrical, 0,7-1, 2 mm long; apex obtuse to rounded;
limbidia weak, generally only a few elongated cells on vaginant and dorsal laminae;
laminal cells hexagonal, 10-12 pm long 16. F. microandrogynus
15 Laminal cells papillose or mammillose:
18 Laminal cells sharply mammillose or with single spinose papilla:
19 Limbidia restricted to proximal $ of vaginant laminae of perichaetial and subperichaetial
leaves, broad at insertion, absent on lower leaves; costa subpercurrent ; seta 0,6-0, 8
mm long 15. F. scleromitrius
19 Limbidia present on most leaves, reaching apex of vaginant laminae; costa short-excurrent;
seta 2-4 mm long 14. F. submarginatus
18 Laminal cells multipapillose, papillae low and blunt:
20 Leaves 0,4-0, 7 mm long; costa ending 6-8 cells below apex 21. F. subobtusatus
20 Leaves 0,8-1 ,4 mm long; costa percurrent or ending just below apex:
21 Laminal cells with 4-6 low, simple papillae over lumen, not obscuring cells, occasionally
almost smooth; terricolous 20. F. erosulus
21 Laminal cells obscured by numerous, low, dense, peripheral papillae; corticolous. .
19. F. bor genii
6 Leaves without limbidia, marginal cells undifferentiated, occasionally bistratose:
22 Plants small; leaves 0,6-0, 9 mm long; laminal cells weakly bulging to sharply mammillose (Sec-
tion Crenularia) 22. F. pseudoserralus
Fissidentaceae
41
22 Plants larger; leaves 1-7 mm long:
23 Plants 5-15 mm long; terricolous or saxicolous, occasionally semi-aquatic; leaves crowded:
24 Leaves unistratose throughout:
25 Dorsal lamina ending above insertion ; stems without axillary hyaline nodules ; costa ending
below apex to percurrent (Section Serridium) :
26 Leaf apices rounded to obtuse; costa ending below apex 26. F. asplenioides
26 Leaf apices acute; costa percurrent 25. F. plumosus
25 Dorsal lamina rounded at insertion, generally crumpled against stem; stems with axillary
hyaline nodules; costa short-excurrent (Section Crispidium) 24. F. glaucescens
24 Leaves partly bistratose:
27 Leaf margins bistratose, rest of lamina unistratose (Section Aloma) 23. F. nitens
27 Superior and dorsal laminae bistratose (Subgenus Pachyfissidens) 27. F. fasciculatus
23 Plants 50-100 mm long, aquatic, floating; leaves distant (Subgenus Octodiceras) 28. F. fontanus
1. Subgenus Aneuron
Aneuron Kindb., Eur. N. Am. Bryin. 2; 165 (1897).
Plants small to minute, light green to glaucous; terricolous. Stems without central strand.
Leaves limbate; costa absent; laminal cells lax.
Sporophyte terminal; seta long; capsules short-cylindrical; exothecial cells thickened in
corners; stomata present in base; spores small.
1. Fissidens enervis Sim in Trans. R. Soc.
S. Afr. 15: 187 (1926). Type: Natal, Pieter-
maritzburg, Town Bush Valley, Sim 9899
(PRE, lecto.!, selected here).
Plants very small to minute, scattered,
light green; terricolous. Stems to 2 mm tall,
simple; in section subround, central strand
absent, cortical cells large, in 2 rows, weakly
thickened. Leaves distant, longer above, very
small below, weakly contorted dry, erect-
spreading wet ; narrowly lingulate to ligulate,
0,8- 1,2 mm long; apex acute, abruptly api-
culate; margins entire, limbidia distinct,
present on all laminae; costa absent. Vaginant
laminae equal, very small, narrow, ending
near mid-leaf; dorsal lamina slightly tapered
proximally, broadly attached at insertion, not
decurrent. Laminal cells elongate-hexagonal,
30-50 //m long, 20-25 nm wide, lax, flat,
smooth.
Perichaetia terminal, leaves narrow, 1 , 2
mm long. Seta to 3 mm long, yellow; capsule
erect, urn ovate to cylindrical, 2, 5-3, 5 mm
long, yellowish; exothecial cells quadrate,
corners thickened ; peristome reddish, deeply
cleft; operculum rostrate, 0,3 mm long;
calyptra just covering operculum, 0,3 mm
long; spores round, 1 2—14 /tm, spiculate. Fig.
Map 7. — • Fissidens erosulus
x Fissidens enervis
Recently reported from central and eastern
Africa (Bizot & Poes, 1979). In Southern Africa, F.
enervis is known only from the type locality in
southern Natal. Map 7.
Vouchers: Van der Bijl PRE-CH5068; Sim 9900.
The species is very distinctive. The large cells,
strong limbidia and absence of a costa will identify
the species.
Fissidentaceae
43
2. Fissidens usambaricus Broth, in Bot.
Jb. 20: 182 (1894); Broth, in Naturl. PflFam.
10: 145 (1924). Type: Tanzania, Usambara,
Lutindi, Holst 3472 (H, holo.).
Plants small, scattered, light green; terri-
colous. Stems 2-3 mm high, simple; in section
elliptical, central strand absent, cortical cells
lax, in 2-3 rows. Leaves distant, little altered
dry; spathulate to Ungulate, 1,2-1 ,4 mm
long; apex obtuse; margins entire, very
weakly bordered by smaller, narrower cells;
costa absent. Vaginant laminae very narrow,
extending of leaf length, termination of
one blade marginal to submarginal; dorsal
lamina gradually tapering proximally, broad-
ly attached to stem, occasionally minutely
decurrent. Laminal cells hexagonal to sub-
hexagonal, 20-37 /tm wide, lax; marginal
cells narrower, short-rectangular; basal cells
of vaginant laminae elongate-rectangular to
rectangular, lax.
Dioicous. Perichaetia terminal; leaves
slightly larger, spathulate. Seta 2-3 mm long,
yellowish; capsule ± erect, urn short-ellip-
tical, 0,4-0, 6 mm long, yellowish; exothecial
cells quadrate, corners strongly thickened;
peristome teeth deeply cleft, red-brown; oper-
culum rostrate, 0,4 mm long; calyptra small,
just covering operculum; spores round, 10-12
fim, weakly papillose. Fig. 8: 8-13.
Map 8. — • Fissidens fontanus
x Fissidens usambaricus
Fissidens usambaricus is known from eastern,
western and Southern Africa. In the Flora area, the
species is found in forests and moist woodlands of
the eastern Transvaal and Zululand. Map 8.
Vouchers: Crosby & Crosby 7801; Vorster 1569.
The elimbate, spathulate leaves, small, isodia-
metric leaf cells and absence of a costa will place
specimens of F. usambaricus. The species has only
recently been reported from Southern Africa (Magill &
Schelpe, 1979).
2. Subgenus Fissidens
Plants small to large, light green or yellow-green to dark green; terricolous or corticolous.
Stems with central strand. Leaves limbate or elimbate; costate; laminal cells generally small,
rarely large, lax.
Sporophyte terminal or lateral; seta long or short; capsule short-cylindrical, erect to
nodding; spores small.
1 . Section Areofissidens
Areofissidens C. Mull., Syn. Muse. 1: 46 (1848).
Plants small, light green to yellow-green; terricolous or rarely corticolous. Leaves limbate
or elimbate; costa ending well below apex. Laminal cells large, lax.
Fig. 8. — Fissidens enervis (1-7): 1. habit, x 1 ; 2. habit, x 10; 3. stem in cross section, x 300; 4. leaf, x40;
5. lower margin of dorsal lamina, x435; 6. leaf apex, x435; 7. capsule and operculum, x20. F. usambaricus
(8-13): 8. habit, xl; 9. habit, x 10; 10. leaf, x25; 11. leaf base, xl70; 12. leaf apex, xl70; 13. capsule and
operculum, x20. F. wageri (14-20): 14. habit, xl; 15. habit, xlO; 16. leaf, x 25; 17. cells at base of vaginant
lamina, x 160; 18. cells at termination of vaginant lamina, x 160; 19. leaf apex, x 160; 20. leaf in median cross
section, xl60. F. splachnifolius (21-27): 21. habit, xl; 22. habit, xlO; 23. leaves, x40; 24. cells at base of
dorsal lamina, x 435; 25. cells at termination of vaginant lamina, x 435; 26. leaf apex, x 435; 27. capsule, x20.
(1-7, Sim 9899; 8-10, Crosby & Crosby 7801; 11-13, Vorster 1569; 14-20, Wager 11 (A \ 21-27, Ecklon s.n.).
44
Fissidentaceae
3. Fissidens wageri Dix. ex Wager in
Trans. R. Soc. S. Afr. 4: 1 (1914); Broth, in
Naturl. PflFam. 10: 145 (1924); Sim, Bryo. S.
Afr. 187 (1926). Type: Natal, Umkomaas,
Wager s.n. (PRE!).
Plants medium-sized, scattered or in
small groups, yellow-green; terricolous. Stems
to 2 mm long, simple; in section round, cen-
tral strand weak, cells with very thin walls,
cortical cells large, thickened, in 4-5 rows.
Leaves crowded above, little altered dry;
elliptical to lingulate, 2-4 mm long; apex
acute to broadly acute; margins elimbate,
entire; costa present, extending to mid-leaf,
ending a few cells above departure from
vaginant laminae; in section cells differen-
tiated. Vaginant laminae equal, extending
of leaf length; dorsal lamina gradually
tapering from mid-leaf to insection. Laminal
cells fusiform to elongate-hexagonal, 60-110
pm long, 20-25 pm wide, flat, smooth.
Autoicous. Perigonia gemmate, axillary.
Perichaetia terminal; leaves undifferentiated,
4 mm long. Seta to 10 mm long, yellow;
capsule nodding to inclined, urn curved, 1 , 5
mm long, yellowish; exothecial cells quadrate,
thin-walled; peristome teeth cleft above
middle, reddish; operculum short-rostrate;
spores round, 12-15 //m, spiculate. Fig. 8:
14-20.
Endemic to Southern Africa, F. wageri is found
in the coastal and inland forests of Zululand, Natal
and Transkei. Map 9.
Vouchers: Pegler 1358; Van der Plank 109.
Map 9. — • Fissidens wageri
X Fissidens hoeegii
Fissidens wageri was originally described as
ecostate. The routine mounting of specimens in
Hoyer’s medium revealed a weak costa in leaves of
the type. Transverse sections of the leaves exposed a
costa with differentiated internal anatomy. The
presence of the costa necessitates moving the species
from subgen. Aneuron to subgen. Fissidens section
Areofissidens.
The species is defined by the large fusiform leaf
cells, weak limbidia, and costa ending far below
apex. The species is not likely to be confused with
any other Southern African species.
4. Fissidens splachnifolius Hornsch. in
Linnaea 15: 145 (1841); Broth, in Naturl.
PflFam. 10: 148 (1924). Type: Cape, Table
Mountain, Ecklon s.n., Aug. 1827 (H-BR!).
Fissidens arnellii P. Varde in Revue bryol. lichen.
23: 266 (1954). Type: Cape, Orange Kloof, Arnell
2180 (PC, holo. !).
Plants very small, scattered, light green;
terricolous. Stems 1-2 mm high, simple; in
section subround, central strand present,
collapsed, cortical cells in 3 rows, large,
irregularly shaped. Leaves erect dry, erect-
spreading wet; asymmetrically ovate-spathu-
late, weakly panduriform, 0,5-1 ,0 mm long;
apex acute, apiculate; margins entire, weakly
bordered by smaller marginal cells; costa
weak, ending well below apex, to i or i of leaf
length. Vaginant laminae to i of leaf length,
termination of one blade laminal to submar-
ginal; dorsal lamina gradually tapering
proximally, weakly decurrent. Upper laminal
cells hexagonal to elongate-hexagonal, 30-47
pm long, 20-25 pm wide, lax, smooth ; margi-
Map 10. — • Fissidens asplenioides
X Fissidens splachnifolius
Fissidentaceae
45
nal cells narrow, rectangular; basal cells
rectangular.
Dioicous. Perichaetia terminal; leaves
undifferentiated, 1,5-1, 8 mm long. Seta 4-5
mm long, reddish yellow; capsule erect to
horizontal, urn very short-cylindrical, weakly
curved, 0,3-0, 6 mm long, reddish yellow;
exothecial cells quadrate to short-rectangular,
longitudinal walls thickened; peristome teeth
deeply cleft, reddish; operculum not seen;
spores round, 17-20 /j. m , weakly papillose.
Fig. 8:21-27.
Endemic to the southwestern Cape, the species
has been collected in the fynbos of Table Mountain
and on the Cape Peninsula. Map 10.
Voucher: Schelpe 7320 (BOL).
Examination of type material clearly indicates
that Sim’s (1926) interpretation of this species was
incorrect. The species described and illustrated by
him is clearly F. pygmaeus ( = F. bifrons)', see note
under that species.
The type specimen of F. splachnifolius is smaller
than plants of more recent collections, both in overall
size and the number of cells across width of upper
leaf. Comparison of a number of other characters
indicate, at this time, that all of these specimens
represent a single species.
5. Fissidens capriviensis Magill, sp. nov.,
F. bocarangensi P. Varde et F. flaccido Mitt,
similis, sed habitatione corticola, statura
minore, forma foliorum et cellulis laminae
minoribus dijfert; in Sectione Areofissidente
Fto. 9. — Fissidens capriviensis (1-7) : 1. habit, x 1 ; 2. habit, X 10; 3. stem in cross section, x 240; 4. leaves,
x60; 5. upper laminal cells, x640; 6. cells at margin of dorsal lamina, x640; 7. leaf apex, x640. F. hoeegii
(8-14): 8. habit, xl; 9. habit, xlO; 10. leaf, x40; 11. leaf base, x435; 12. cells at margin of dorsal lamina,
X640; 13. leaf apex, x435; 14. capsule, x20. (1-7, Vahrmeijer 122; 8-14, Magill 5584).
46
Fissidentaceae
propter limbidias bene distinctas laminis omni-
bus, cellulas laminae magnas, costam dis-
tinctam terminatam infra apicem ponitur.
Type: South West Africa/Namibia, Ca-
privi Strip, on trees in mixed dry woodlands ,
Vahrmeijer 122 (PRE, holo.; MO; NY).
Plants small, scattered, dark green; cor-
ticolous. Stems 1-2 mm high; in section oval,
central strand very small or absent, cortical
cells in 3-4 rows, large, thin-walled, smaller
toward margin. Leaves distant below,
crowded above, bent backwards dry, erect-
spreading wet; elliptical to spathulate,
0,6-1 , 1 mm long, smaller below; apex acute;
base narrowed; margins entire, limbate on all
laminae; costa weak, ending well below apex.
Vaginant laminae equal, of leaf length;
dorsal lamina gradually tapering proximally,
rounded at insertion. Laminal cells elongate-
hexagonal, 20-32 pm long, 5-10 pm wide,
thin-walled, flat; basal cells rectangular,
25-37(45) /mi long, 12-15 pm wide, thin-
walled.
Sporophyte unknown. Fig. 9: 1-7.
The plants were collected in woodlands of the
Caprivi Strip, northeastern South West Africa/
Namibia. Map 11.
Map 11. — • Fissidens curvatus
x Fissidens capriviensis
Voucher: Type only.
The species is similar to F. bocarangensis P.
Varde and F. flaccidus Mitt, but differs from both in
being corticolous, as well as in the smaller plant size,
leaf shape and shorter leaf cells. The well defined
limbidia, large leaf cells and distinct costa distinguish
the species from other Southern African species.
2. Section Fissidens
Fissidens emend. Norkett ex Gangulee, Moss. E. India 453 (1971).
Plants small to medium; terricolous. Leaves with limbidia on all laminae, strong or weak;
costa percurrent to short-excurrent. Laminal cells small, smooth, flat or mammillose.
6. Fissidens bryoides Hedw., Spec. Muse.
153 (1801); Broth, in Naturl. PflFam. 10: 146
(1924); Crum, Mosses of the Great Lakes
Forest 59 (1973); Pursell & Hoe in J. Hattori
bot. Lab. 43: 83 (1977). Type: Europe.
Fissidens androgynus Bruch ex Krauss in Flora,
Jena 29: 134(1846); Broth, in Naturl. PflFam. 10: 146
(1924); Sim, Bryo. S. Afr. 197 (1926). Type: Cape,
Devil’s Peak, Krauss s.n. (BM !).
? Fissidens remolifolius C. Mull., Syn. Muse. 1 : 60
(1848); Broth, in Naturl. PflFam. 10: 146 (1924). Type
Cape, Zeyher 41.
Fissidens gueinzii C. Mull, in Linnaea 37: 168
(1872); Broth, in Naturl. PflFam. 10: 146 (1924);
Sim, Bryo. S. Afr. 198 (1926). Type: Cape, Gueinzius
s.n. (BM!).
Fissidens subremotifolius C. Mull, in Hedwigia 38:
54 (1899); Broth, in Naturl. PflFam. 10: 146 (1924).
Type: Transvaal, Lydenburg, Wilms s.n., Apr. 1887,
herb. Jack (G, holo.!).
Fissidens malacobryoides C. Mull, in Hedwigia
38: 55 (1899); Broth, in Naturl. PflFam. 10: 146
(1924); Sim, Bryo. S. Afr. 200 (1926). Type: Cape,
Somerset East, Boschberg, MacOwan s.n. (G!).
Plants medium-sized forming loose
groups, dark green; terricolous. Stems to 10
mm high, infrequently branched below; in
section subround to elliptical, central strand
present, inner cortical cells large, thin-walled,
outer cortical cells in 1-2 rows, smaller,
incrassate. Leaves evenly spaced, contorted
dry, erect-spreading wet; ovate or ligulate to
elliptical, occasionally asymmetrically ovate-
elliptical, 1-2 mm long; apex broadly acute
to cuspidate; margins entire to serrulate at
apex; limbidia strong on all laminae, con-
fluent with costa or ending a few cells below
apex, generally ending above base of dorsal
laminae; costa percurrent to short-excurrent.
Fissidentaceae
47
Vaginant laminae equal, to $- of leaf length;
dorsal lamina abruptly narrowing proximally,
rounded to insertion, occasionally weakly
decurrent. Laminal cells rounded, quadrate to
hexagonal, 8-12 //m long, flat or occasionally
bulging, smooth.
Autoicous. Perichaetia terminal; leaves
somewhat larger. Seta 3-6 mm long,
yellowish; capsules erect to inclined, urn
straight to curved, (0,6-)0, 8-1,0 mm long,
reddish yellow; exothecial cells short-rectan-
gular, longitudinal walls thickened, cells at
mouth quadrate; peristome deeply cleft,
reddish; operculum rostrate, 0,4-0, 5 mm
long; spores round, 11-21 //m, weakly
papillose. Fig. 10: 1-8.
This widely distributed species is known from
Europe, Africa, North America, southern South
America, Asia and Oceania. In Southern Africa the
species is found in grasslands of the Transvaal,
Orange Free State, Lesotho, Natal and the eastern,
central and southwestern Cape. Sporophytes were
only found on specimens at upper elevations in and
around Lesotho. Map 12.
Vouchers: Crosby & Crosby 7771; Garside 6201;
Mag ill 3328, 4367; Schelpe 21 10.
Fissidens bryoides has been confused with several
other species of section Fissidens. It is recognized by
the vaginant laminae being equal and the large, flat
leaf cells.
The species varies in development of both costa
and limbidium. The costa is generally percurrent to
short-excurrent, but may end 1-3 cells below apex.
The limbidium may be very strong, extending from
insertion to apex where it is confluent with the costa
or it may be weak, ending above insertion and below
apex.
7. Fissidens hoeegii P. Varde in K. norske
Vidensk. Selsk. Skr. 5: 130 (1932). Syntypes:
Natal, Umgeni, Pietermaritzburg, H<\>eg 423,
430 (PC!), 432 (PC!; PRE!).
Plants small, in loose groups, yellow-
green to dark green; terricolous. Stems 3-6
mm tall, simple; in section subround, central
strand small, inner cortical cells large, in two
rows, lax, smaller toward margin, outer
cortical cells stereids, in 1-2 rows, brownish.
Leaves not crowded, contorted dry, erect-
spreading wet; ligulate to oblong-elliptical,
1 ,2-1 ,3 mm long; apex acute; margins entire,
limbidia present on all laminae, ending just
below apex, and ending above insertion on
dorsal lamina, broader on proximal vaginant
laminae, to 4-6 rows of rectangular cells;
costa percurrent. Vaginant laminae equal, to |
of leaf length; dorsal lamina gradually
tapering proximally, extending to insertion or
slightly decurrent. Laminal cells hexagonal to
subhexagonal, 5-7 //m long, superficially
bulging, smooth; quadrate in proximal vagi-
nant laminae.
Perichaetia terminal; leaves ligulate, to
1 , 5 mm long. Seta 4 mm long, reddish yellow ;
capsule ± erect, urn cylindrical, 0 , 8 mm long,
yellowish; exothecial cells short-rectangular,
longitudinal walls thickened, quadrate at
mouth, strongly incrassate ; peristome cleft to
near base, yellowish red; operculum short-
rostrate ; spores round, 25-32 /an, very weakly
papillose. Fig. 9: 8-14.
Recently reported from Tanzania (Bizot & Poes,
1979). In Southern Africa, F. hoeegii is known from
kloof forests of Natal and eastern Orange Free State.
Map 9.
Vouchers: Cholnoky 104; Magill 5584.
The species is similar to F. bryoides but differs
in smaller, bulging leaf cells and a broad limbidium
on proximal vaginant laminae. The equal vaginant
laminae will separate F. hoeegii from the other species
in this section.
8. Fissidens rufescens Hornsch. in Lin-
naea 15: 153 (1841); Broth, in Naturl. PflFam.
10: 146 (1924); Sim, Bryo. S. Afr. 198 (1926).
Syntypes: Cape, Lion’s back, Ecklon s.n..
Sept. & Nov.; Table Mountain, Ecklon s.n.,
Oct.; Devil’s Peak, Ecklon s.n., Dec. (All
BM!).
Fissidens megalotis Schimp. ex C. Miill. in Bot.
Ztg 16: 154 (1858); Broth, in Naturl. PflFam. 10:
149 (1924); Sim, Bryo. S. Afr. 193 (1926). Type:
Cape, Greenkloof, Breutel s.n. (BM, holo.!).
48
Fissidentaceae
Fissidens breutelii, p.p. quoad Cape, Soutkloof,
Breutel s.n. (BM!), syntype of F. breutelii Schimp. ex
C. Mull, in Bot. Ztg 17: 198 (1859).
Fissidens macowanianus C. Miill. in Hedwigia 38:
58 (1899); Broth, in Natiirl. PflFam. 10: 146 (1924).
Type: Cape, Somerset East, Boschberg, MacOwan
s.n., 1883 (HBG ! ; BM!).
Fissidens minutiretis Dix. in K. norske Vidensk.
Selsk. Skr. 1932 (4): 5 (1932). Type: Cape, Signal
Hill, Moss & Moss 9261 (BM, holo.!).
Plants small to medium, scattered or
forming loose groups, dark green to yellow-
green; terricolous. Stems 4-8 mm high,
simple; in section elliptical, central strand
present, inner cortical cells large, outer
cortical cells in 2 rows, stereids to substereids.
Leaves ± crowded, compressed or weakly
contorted dry, erect-spreading wet ; ligulate to
elliptical or frequently asymmetrical, broadly
ovate-ligulate 0,5-1, 8 mm long; apex acute,
cuspidate; margins serrulate at apex, entire
below or occasionally serrulate on vaginant
laminae, limbidia present on all laminae,
strong or weak, confluent with costa or ending
a few cells below apex and ending above inser-
tion on dorsal lamina, submarginal to inter-
marginal by 1-8 cells, on proximal vaginant
laminae, occassionally spurred; costa short-
excurrent. Vaginant laminae open to closed,
to f of leaf length; termination of one blade
submarginal to laminal or almost at costa;
dorsal lamina gradually tapering proximally,.
just reaching insertion or weakly decurrent,
rarely rounded to insertion. Laminal cells
rounded, quadrate to subhexagonal, 4-6 jum
long, weakly thickened, mammillose to
bulging, smooth.
Perichaetia terminal; leaves elongate,
asymmetrically ovate-ligulate, 2,0-2, 5 mm
long. Seta 5-7 mm long, yellowish; capsule
inclined to horizontal, urn ± asymmetrical,
curved, 0,8-1 ,0 mm long, yellow-brown;
exothecial cells quadrate, longitudinal walls
thickened; peristome deeply cleft, reddish;
operculum rostrate; spores round, 15-18 //m,
weakly papillose. Fig. 10: 9-16.
Endemic to Africa, F. rufescens has been reported
from the eastern and southern parts of the continent.
In Southern Africa the species is widespread in drier
regions of Transvaal, Natal, Lesotho, Orange Free
State and Cape. A few specimens have also been
collected in Botswana. The species is not known
fiom South West Africa/Namibia, although its
presence in the northwestern Cape indicates that it
may be found in the southern parts of South West
Africa/Namibia. Map 13.
Vouchers: De Winter 9601; Magill 3204, 4648,
6250; Smook 1412.
The species is recognized by the unequal vagi-
nant laminae, intermarginal limbidia on proximal
vaginant laminae and small, mammillose leaf cells.
The character of intermarginal limbidia was found
to be variable but useful. In the western Cape the
limbidia are generally 6-8 cells deep on the proxi-
mal vaginant laminae. In other parts of the Cape,
Orange Free State and Natal the depth is most
frequently 3-6 cells, while in the Transvaal 1-3 is
more common. Another character, that of open
vaginant laminae, also shows similar variation. In the
western Cape the laminae are conspicuously open,
wet or dry, while eastern and northern collections
have laminae only weakly open to closed.
The specimens reported from Southern Africa
(Schelpe, 1979) as F. vittatus Hook. f. & Wils. belong
here. Judging from the description and illustrations
published by Scott & Stone (1976), it is probable
Fig. 10. — Fissidens bryoides (1-8): 1. habit, X 1 ; 2. habit, X 10; 3. leaf, X 35; 4. leaf base, x435; 5. cells at
termination of vaginant lamina, x435; 6. leaf apex, x435; 7. upper laminal cells, x640 ; 8. capsule, x20. F.
rufescens (9-16): 9. habit, x 1 ; 10. habit, X 10; 11-13. leaves, X35; 14. leaf base, x435; 15. cells at margin of
dorsal lamina, x 640; 16. leaf apex, x435. F. marginatus (17-24): 17. habit, x 1 ; 18. habit, x 10; 19-20. leaves,
x 35; 21. cells at lower margin of dorsal lamina, x435; 22. cells at lower margin of vaginant lamina, x435;
23. cells at termination of vaginant lamina, x435; 24. leaf apex, x435. F. simii (25-31): 25. habit, xl; 26.
habit, x 10; 27. leaf, x 55; 28. leaf base, x 170; 29. cells at termination of vaginant lamina, X 170; 30. cells at
margin of dorsal lamina, X640; 31. leaf apex, x 170. (1-2 & 8, Magill 4614; 3-7, Bosman 1165; 9-10, Schelpe
4917; 11-16, Magill 4076; 17-18, Wager 294; 19-24, Rehmann 290; 25-26, Oliver 7148; 27-31, Sim 9907).
Fissidentaceae
49
50
Fissidentaceae
that the Australian species is synonymous with F.
rufescens; however, further observations must be
made.
9. Fissidens marginatus Schimp. ex C.
Mull, in Bot. Ztg 16: 154 (1858); Broth, in
Naturl. PflFam. 10: 146 (1924); Sim, Bryo. S.
Afr. 199 (1926). Type: Cape, Table Mountain,
Ecklon s.n., (BM, holo.!).
Fissidens breutelii Schimp. ex C. Mull, in Bot.
Ztg 17: 198 (1859). Type: Cape, Gnadenthal, Kuhn
s.n. (BM, lecto. !, selected here).
Fissidens ischyrobryoides C. Mull, in Hedwigia 38 :
55 (1899); Broth, in Naturl. PflFam. 10: 146 (1924).
Type: Cape, Devil’s Peak, Rehmann 290 (PRE!).
Plants small to medium, forming loose
groups, green, glossy; terricolous. Stems 4-8
mm high, simple; in section subround, central
strand present, inner cortical cells large, outer
cortical cells in 1-3 rows, smaller, incrassate.
Leaves evenly spaced, weakly contorted dry,
widespreading wet; ligulate to ovate-lingulate
or elliptical, 1-2 mm long; apex acute to
broadly acute, cuspidate; margins serrulate to
serrate at apex, entire below, limbidia strong
on all laminae, confluent with costa or ending
just below apex; costa short-excurrent.
Vaginant laminae -J— £ of leaf length, one blade
abruptly rounded distally, terminating on
costa; dorsal lamina only slightly tapering
proximally, weakly decurrent. Laminal cells
irregularly shaped, rounded, quadrate to hex-
agonal or angular, 8-12 pm long, weakly
thickened, bulging.
Perichaetia terminal, occasionally poly-
setaceous; leaves asymmetrical ovate-lingu-
late, 1 ,4 mm long. Seta 5-7 mm long, yellow
to reddish; capsule inclined to horizontal, urn
asymmetrical, to 1 mm long, reddish; exothe-
cial cells short-rectangular, longitudinal walls
thickened; peristome deeply cleft, reddish;
operculum long-rostrate, 0,6 mm long; spores
round to subreniform, 1 5—20 pm, smooth,
green. Fig. 10: 17-24.
Recently reported from Tanzania by Bizot,
Poes & Sharp (1979). In Southern Africa, F. margi-
natus is found in drier coastal regions of the western
and southern Cape. Map 14.
Vouchers: Esterhuysen 24364; Pare 4; Stokoe
PRE-CHI 281 3.
The abrupt distal rounding of one blade of the
vaginant laminae and its strong termination on the
costa will separate F. marginatus from all other
Southern African species except F. simii. The size
and shape of leaves and laminal cells will separate
the latter.
Map 14. — • Fissidens marginatus
x Fissidens scleromitrius
The syntypes of F. breutelii Schimp. ex C. Mull,
are discordant elements; the Kuhn specimen is
selected above as lectotype and the Breutel specimen
is treated in the synonomy of F. rufescens Hornsch.
10. Fissidens simii Schelpe in Mem. bot.
Surv. S. Afr. 43: 5 (1979). Syntypes: Natal,
Pietermaritzburg, Sim 9903, 9907, 9909
(PRE!).
Fissidens aristatus Sim, horn, illeg., Bryo. S.
Afr. 200 (1926), non Broth. (1916).
Plants small, scattered, light green; terri-
colous. Stems 3-5 mm high, simple; in section
subround, central strand present, cortical
cells smaller toward margin, outer row sub-
stereids, reddish. Leaves distant, little altered
dry, erect-spreading wet; elliptical to lanceo-
late, 0,8-1 , 1 mm long; apex short-acuminate;
margins entire, strongly limbate throughout;
costa percurrent to short-excurrent. Vaginant
laminae to mid-leaf, one blade narrowing
distally, terminating on costa; dorsal lamina
rounded at insertion. Laminal cells quadrate
to rhombic, 12-16 (-20) //m long, weakly
incrassate, smooth, flat.
Sporophyte not seen. Described by Sim
(1926) as terminal, seta long; capsule inclined
to horizontal, urn short-cylindrical; oper-
culum short-rostrate. Fig. 10: 25-31.
Fissidens simii was described from Natal. The
species, although rare, also occurs in the eastern
Cape, eastern Transvaal and in eastern Africa
(Bizot & Poes, 1979). Map 15.
Vouchers: Oliver 7148; Wager PRE-CH11966.
Fissidentaceae
51
Map 15. — • Fissidens borgenii
X Fissidens simii
The species is identified by its elliptical to lanceo-
late leaves, termination of one blade of the vaginant
laminae on the costa and large, flat laminal cells.
11. Fissidens curvatus Hornsch. in Lin-
naea 15: 148 (1841); Broth, in Natiirl. PflFam.
10: 146 (1924); Sim, Bryo. S. Afr. 188 (1926).
Type: Cape, Mr Auret’s garden, Lowen-
riicken, Ecklon s.n., 24 Oct. 1827 (H-BR!).
Fissidens laxifolius Hornsch. in Linnaea 15: 149
(1841); Broth, in Natiirl. PflFam. 10: 149 (1924);
Sim, Bryo. S. Afr. 189 (1926). Syntypes: Cape, Mr
Breda’s garden under Table Mountain, Ecklon s.n.,
15 Aug. 1827; Table Mountain, Ecklon s.n., 1 Jun.
1827; Table Mountain, Ecklon s.n., 30 Aug. 1826
(H-BR!).
Fissidens cuspidatus C. Mull, in Linnaea 17: 588
(1843); Broth, in Natiirl. PflFam. 10: 146 (1924); Sim,
Bryo. S. Afr. 196 (1926). Type: Cape, Krakakammae,
Ecklon s.n. (Hb. Kunze).
Conomitrium gracile Hampe ex C. Mull, in Bot.
Ztg 17: 197 (1859). Fissidens gracilis (Hampe) Jaeg.,
horn, illeg., Enum. Fiss. 12 (1869), vide Dix. in Sim,
Bryo. S. Afr. 197 (1926). Type: Cape, Elim, Breutel
s.n.
Fissidens pycnophyllus C. Mull, in Hedwigia 38:
57 (1899); Broth, in Natiirl. PflFam. 10: 147 (1924);
Sim, Bryo. S. Afr. 187 (1926). Type: Cape, Cape
Town, Rehmann 293 (PRE!).
Plants small, scattered, yellow-green,
glossy; terricolous. Stems 1-3 mm long,
simple; in section subround, central strand
present, inner cortical cells large, in single
row, outer cortical cells in 2 rows, small,
substereids. Leaves larger above, slightly
contorted dry, erect-spreading wet; linear-
lanceolate to lanceolate, 0,7-0, 8 mm long;
apex acute; margins entire, strongly limbate
throughout; costa percurrent. Vaginant lami-
nae narrow, to upper | of leaf, termination
of one blade variable, marginal or laminal;
dorsal lamina gradually tapering proximally,
generally ending above insertion. Upper
laminal cells irregularly shaped, rounded,
angular, elongate-hexagonal or rectangular,
10-17 pm long, smooth, flat; basal cells of
vaginant laminae generally rectangular.
Autoicous. Perigonia gemmate, axillary.
Perichaetia terminal; leaves asymmetrically
broadly ovate-lanceolate, 1,2 mm long. Seta
(1 ,2-)4-6 mm long, yellow; capsule inclined,
urn short-cylindrical, 0,8 mm long, yellowish;
exothecial cells quadrate to short-rectangular,
weakly thickened; peristome deeply cleft,
reddish; operculum rostrate; spores round,
14-17 pm, weakly papillose. Fig. 11: 1-8.
Endemic to Southern Africa, F. curvatus is
infrequently collected in grasslands or shrublands
of the southwestern, central, southern and eastern
Cape Province, Orange Free State, Natal, Swaziland
and the central, eastern and northern Transvaal.
Map 11.
Vouchers: Kemp 853; Magill 3291, 4087;
Vahrmeijer PRE-CHI 2644.
The small, narrow leaves and very strong yel-
lowish border distinguish specimens of F. curvatus
(see note under F. stellenboschianus). The shape of the
laminal cells is rather variable. Individual collections
generally exhibit only one of the cell patterns known
for the species, but several collections have been seen
with a heterogeneous cell pattern.
Seta length and degree of inclination of the
capsule also show considerable variation. The type
of F. curvatus has a very short seta and slightly inclined
capsule. Frequently specimens have setae up to
6 mm long and capsules nearly horizontal.
12. Fissidens stellenboschianus Dix. ex
Sim, Bryo. S. Afr. 195 (1926). Type: Cape,
Stellenbosch, Wager 612 (BM, holo.!; PRE!).
Plants small, scattered, light green;
terricolous. Stems to 1 mm long, simple; in
section oval, central strand small, cortical
cells large, lax, outer row smaller, incrassate.
Leaves crowded, little altered dry; narrowly
elliptical, 0, 5-0,7 mm long; apex acute;
margins serrulate at apex, entire below,
weakly limbate on all laminae, leaves of sterile
plants with a few elongated cells on vaginant
and dorsal laminae, limbidia stronger on
leaves of fertile plants, extending to just above
termination of vaginant laminae, ending
below apex and above base on dorsal lamina;
costa percurrent to very short-excurrent.
Fissidentaceae
53
Vaginant laminae ±± leaf length, termination
of one blade submarginal ; dorsal lamina
gradually tapering proximally to insertion.
Laminal cells irregularly shaped, shortly
oblong-hexagonal, 12-20 urn long; basal cells
of vaginant laminae short-rectangular.
Dioicous. Perichaetia terminal; leaves
narrow, asymmetrically ovate-lingulate, to 1
mm long. Seta to 3 mm long, yellowish;
capsule ± erect, urn short-cylindrical, 0,5
mm long; exothecial cells quadrate to short-
rectangular, incrassate; peristome irregular,
teeth not cleft, lanceolate, apices abruptly
rounded, 0,1 mm long; operculum short-
rostrate; spores not seen. Fig. 11 : 18-23.
Fissidens stellenboschianus has recently been
reported from Tanzania (Bizot, Poes & Sharp, 1979).
In Southern Africa the species is known only from the
fynbos biome of the southwestern Cape. Map 16.
Voucher: Type only.
Fissidens stellenboschianus is very closely related
to F. curvatus. The two species are separated primarily
on the basis of limbidial development. The limbidia of
F. curvatus are strong on all laminae. Extending from
above the insertion to the apices, the limbidia are
confluent with the costae. The limbidia of F. stellen-
ET~—| I.T 1 T 1 T— i — T~ — ’ T
1 1 | |
-
iirnffi: xl
44ffi4Ti — ~n~
—f
7,
— HH— r-V4-41LJ_(
® r K 1 — 1 — i — Lj
rrmm£ L
.i
-444 IpflTt
+TTTT \ lil
T]xr i
rTr^TITTr
1M V -
rmtma
Map 1 6. — • Fissidens submarginatus
x Fissidens stellenboschianus
boschianus are weak to absent in leaves of sterile
plants and only weakly developed on vegetative
leaves of fertile plants. The limbidia are stronger on
perichaetial and subperichaetial leaves, but do not
reach the apex. Fissidens stellenboschianus is prob-
ably only a variation of F. curvatus; however, inter-
mediates were not seen. The species is provisionally
maintained until additional material can be examined.
13. Fissidens aciphyllus Dix. in Trans. R.
Soc. S. Afr. 18: 250 (1930). Type: Transkei,
Port St Johns, Wager 927 (BM, holo. ! ; PRE!).
Plants small, scattered, light green; terri-
colous. Stems 3-4 mm tall, simple; in section
subround, central strand present, cortical
cells lax, outer row smaller, incrassate. Leaves
larger above, little altered dry ; ± asymmetri-
cal, ovate-lanceolate to lanceolate or narrowly
elliptical, 1, 0-2,0 mm long; apex short-
acuminate; margins entire, limbate on all
laminae, broader on vaginant lamina; costa
percurrent to short-excurrent. Vaginant lami-
nae extending to mid-leaf, termination of one
blade laminal; dorsal lamina narrowing
proximally to insertion. Laminal cells large,
lax, hexagonal to elongate-hexagonal, 30-45
Map 17. — • Fissidens aciphyllus
x Fissidens nitens
Fig. 11. — Fissidens curvatus (1-8): 1. habit, xl;2. habit, x 10; 3. leaf, x40;4.1eaf, x 70; 5. cell at base of
dorsal lamina, x435; 6. laminal cells, x435; 7. cells at margin of dorsal lamina, x640; 8. leaf apex, x435.
F. aciphyjlus (9-17): 9. habit, xl;10. habit, x 10; 11. leaf, x 33; 12-13. leaves, x 15; 14. cells at base of vagi-
nant lamina, x 170; 15. cells at termination of vaginant lamina, x 170; 16. leaf apex, x 170; 17. laminal cells,
x435. F. stellenboschianus (18-23): 18. habit, x 1 ; 19. habit, x 10; 20. leaf, x 80; 21. leaf base, x435; 22. leaf
apex, x 435; 23. cells at margin of vaginant lamina, x435. F. nitens (24-30): 24. habit, x 1 ; 25. habit, x 10; 26.
stem in cross section, x290; 27. leaf x33; 28. leaf base, x 170; 29. upper laminal cells at margin, x640; 30.
leaf apex, x 170. (1-2, Kemp 853; 3-8, Vahrmeijer PRE-CH12646; 9-17, Wager 1446; 18-23, Wager 612; 24-30,
Rehmann 289).
54
Fissidentaceae
//m long, 12-17 //m wide, flat, smooth; basal
cells of vaginant laminae rectangular.
Perichaetia terminal; leaves undifferen-
tiated. Seta 2,5 mm long, yellowish; capsules
erect, urn short-cylindrical, 0,6 mm long;
exothecial cells rectangular, weakly thickened;
operculum rostrate, 0,4 mm long; capsules
immature, therefore other structures not seen.
Fig. 11: 9-17.
Endemic to Southern Africa, F. aciphyllus has
been collected in forests and woodlands of Transkei,
Natal and the Drakensberg of eastern Orange Free
State. A specimen from the central Transvaal was
collected on rock near water. Map 17.
Vouchers: Sim 9940; Wager 1446; Watson 1582.
Fissidens aciphyllus is identified by its strong
limbidia on all laminae, percurrent to short-excur-
rent costae and very large hexagonal to elongate-
hexagonal leaf cells.
3. Section Semilimbidium
Semilimbidium C. Mull, emend. Norkett ex Gangulee, Moss. E. India 454 (1971).
Section Semilimbidium C. Miill., Gen. Muse. Fr. 60 (1900).
Plants small; terricolous or corticolous. Leaves, or frequently only perichaetial leaves,
with limbidia restricted to vaginant laminae. Laminal cells small, smooth or mammillose or
papillose.
14. Fissidens submarginatus Bruch ex
Krauss in Flora, Jena 29: 133 (1846); Broth,
in Natiirl. PflFam. 10: 149(1924); Sim, Bryo.
S. Afr. 193 (1926). Type: Natal, Krauss s.n.
(BM!).
Fissidens opacifolius Mitt., p.p. in Trans. Finn. Soc.
Fond. 23; 54 (1860); Broth, in Natiirl. PflFam. 10:
149 (1924). Syntypes: Natal, Gueinzius s.n., non
West Africa, Vogel s.n. (both NY!).
Fissidens eschowensis Broth. & Bryhn in Forh.
Vidensk. Selsk. Krist. 1911 (4): 8 (1911); Broth, in
Natiirl. PflFam. 10: 149 (1924). Type: Natal, Zulu-
land, Eshowe, H. Bryhn s.n., Jan. 1909 (H-BR,
holo. !).
Fissidens papillifolius Dix. in Trans. R. Soc. S.
Afr. 8: 187 (1920); Broth, in Natiirl. PflFam. 10: 149
(1924); Sim, Bryo. S. Afr. 194 (1926). Type: Natal,
Albert Falls, Umgeni Nook, Sim 8709 (BM, holo.!;
PRE!).
Fissidens urceolatus Wag. & Dix. ex Sim, Bryo. S.
Afr. 194 (1926). Type: Transvaal, Pretoria, Wager 264
(PRE!).
Plants small, forming groups, green to
yellow-green ; terricolous. Stems 3-5 mm long,
simple; in section round, central strand
present, inner cortical cells large, outer corti-
cal cells in 2 rows, smaller, strongly incrassate.
Leaves larger above, equally spaced, curved
and weakly contorted dry, erect-spreading
wet; ligulate or asymmetrically ovate-ellipti-
cal, 0,8-1 ,2 mm long; apex broadly acute to
obtuse, cuspidate; margins limbate on vagi-
nant laminae, crenulate by projecting cells on
other laminae, limbidia narrow, ending above
insertion, marginal throughout; costa strong,
short-excurrent. Vaginant laminae ± equal, to
3 of leaf length; dorsal lamina tapering
proximally to insertion. Laminal cells
quadrate to hexagonal, 5-7 //m long, super-
ficially bulging, cells sharply mammillose,
rarely with 3-4 low, sharp papillae over
lumen.
Autoicous. Perigonia gemmate, axillary.
Perichaetia terminal ; leaves somewhat larger,
little differentiated. Seta 2-4 mm long,
yellowish; capsule erect to inclined, urn short,
cylindrical to elliptical, 0, 5-1,0 mm long,
yellowish, mouth reddish; exothecial cells
quadrate, strongly thickened in corners;
peristome cleft to lower teeth 0 , 3 mm long,
yellowish red; operculum conical, to 0,7 mm
long; spores round, 12-15 //m, granulate.
Fig. 12: 1-8.
Fissidens submarginatus is found in eastern and
Southern Africa. In the Flora area the species is
infrequently collected in northern, central, southern
and eastern Transvaal, Swaziland, Zululand, Natal
and Transkei. Map 16.
Vouchers: Magill 3429, 5069; Russell 2644;
Schelpe 7523; Smook 1362; Van Vuuren 1737.
The species is identified by the well-developed
limbidia on vaginant laminae of all leaves and
strongly mammillose leaf cells. This species could
easily be confused with sterile specimens of F. sclero-
mitrius. The shorter leaves, termination of limbidia
above insertion and strongly mammillose cells of F.
submarginatus should separate the species.
A narrow leaf form, with bulging leaf cells, has
been collected in the Transkei. The bulging leaf cells
give the plants a different appearance from the more
common sharply mammillose specimens; however
other character differences were minor. The strong
limbidia on vaginant laminae of all leaves should
place these specimens.
Fissidentaceae
55
15. Fissidens scleromitrius ( Besch .) Broth.
in Natiirl. PflFam. 10: 149 (1924). Type:
Madagascar.
Conomitrium scleromitrius Besch. in J. Bot., Paris
5: 143 (1891). Moenkemeyera scleromitria (Besch.)
P. Varde in Arch. Bot. Mem 2: 21 (1930).
Fissidens brevisetus Sim, Bryo. S. Afr. 194 (1926).
Type: Natal, New Hanover, Sim 9906 (PRE, holo.!).
Plants medium-sized, light green to
yellow-green; terricolous. Stems 4-6 mm tall,
simple; in section subround, central strand
present, inner cortical cells thin-walled, outer
cortical cells in 1-2 rows, smaller, incrassate.
Leaves larger above, little contorted dry,
erect-spreading wet; oblong-lanceolate to
asymmetrically ovate-lingulate, 0 , 8-1 , 5 mm
long; apex acute to broadly acute; margins
crenulate by marginal cell papillae, limbidia
on vaginant laminae only, restricted to
proximal $ of laminae, broad at insertion,
strong on perichaetial and subperichaetial
leaves, absent in lower leaves; costa subper-
current, ending 2-3 cells below apex. Vaginant
laminae to \ of leaf length, termination of one
blade laminal; dorsal lamina tapering proxi-
mally or occasionally abruptly rounded
below, ending just above insertion. Upper
laminal cells thin-walled, hexagonal to sub-
hexagonal, 7-10 pm long, superficially
bulging, weakly mammillose; basal cells
quadrate, smooth.
Dioicous. Perichaetia terminal; leaves
lanceolate, to 2,5 mm long. Seta very short,
0,6-0, 8 mm high, yellowish; capsule erect,
urn short-cylindrical, 0,6-0, 7 mm long, wide
at mouth, yellowish, mouth reddish; exothe-
cial cells short-rectangular, thin-walled, at
mouth cells very small, quadrate, in 2-4 rows;
peristome not seen; operculum rostrate, 0,8
mm long; spores round, 12 pm, weakly papil-
lose. Fig. 12: 9-15.
Fissidens scleromitrius is known from the East
African Islands and Natal. The species is rare,
collected in Southern Africa only once by T. R. Sim
near New Hanover. Map 14.
Voucher: Sim 9906.
When fruiting the species is recognized by its
very short seta. The distinct limbidia on proximal
vaginant laminae, weakly mammillose leaf cells and
oblong-lanceolate leaves, to 1 ,5 mm long, characterize
the species.
16. Fissidens microandrogynus Dix. in S.
Afr. J. Sci. 18: 306 (1922); Broth, in Natiirl.
PflFam. 10: 149 (1924). Type: Zimbabwe,
Bulawayo, Wager 895 (BM, holo.!; PRE!).
Fissidens calochiorus Dix. in S. Afr. J. Sci. 18: 306
(1922); Broth, in Natiirl. PflFam. 10: 150 (1924);
Sim, Bryo. S. Afr. 189 (1926). Syntypes: Zimbabwe,
Victoria Falls, Sim 8891 (c.fr.), 8882 (BM!; PRE!).
Fissidens hyalobasis Dix. ex Sim, Bryo. S. Afr. 195
(1926). Type: Transvaal, Moorddrift, Wager 406
(BM, holo.!; PRE!).
Plants small to medium, in loose groups,
dark green to yellow-green ; terricolous. Stems
3-5 mm long, simple; in section subround,
central strand small, inner cortical cells thin-
walled, outer cortical cells in 1-2 rows,
smaller, incrassate. Leaves ± distant, little
altered dry; oblong-elliptical to asymmetri-
cally ovate-lingulate, 0,7-1 ,2 mm long; apex
obtuse to rounded in lower and median leaves,
frequently acute in upper leaves; margins
entire, weakly limbate to elimbate, frequently
with only a few differentiated cells on vaginant
laminae; costa subpercurrent by 2^4 cells.
Vaginant laminae to \ of leaf length, termina-
tion of one blade laminal; dorsal lamina
gradually tapering to insertion, occasionally
abruptly rounded at insertion. Laminal cells
rounded, regularly hexagonal to quadrate,
12-16 pm long, smooth.
Perichaetia terminal; leaves curved,
asymmetrically ovate-ligulate, 1,4-1, 5 mm
long, limbidia on vaginant laminae only. Seta
2,0-2, 5 mm long, yellowish; capsule erect to
inclined, urn short-cylindrical, 0,5-0, 6 mm
long, yellowish; exothecial cells rectangular,
longitudinal walls thickened; peristome deeply
cleft, reddish; operculum rostrate; spores
round, 12-15 pm, weakly papillose. Fig. 12:
16-21.
Map 18.— • Fissidens microandrogynus
X Fissidens pygmaeus
56
Fissidentaceae
Fissidentaceae
57
This species is known from eastern and Southern
Africa. In the Flora area it is infrequently collected
in Botswana, the eastern and central Transvaal, Natal,
Lesotho and the Orange Free State. Map 18.
Vouchers: Bosman 1594; Magill 4749; Potts
PRE-CH5126.
The short, broad leaves, weak limbidia and
smooth leaf cells will help to identify this species.
Fissidens microandrogynus might be confused with
F. parvilimbatus ; larger leaves and smaller leaf cells
should separate the former.
17. Fissidens parvilimbatus Sim, p.p.,
Bryo. S. Afr. 190 (1926). Syntypes: Natal,
New Hanover, Sim 9905; Albert Falls, Sim
9912 (PRE!); non Sim 9915 (PRE!).
Fissidens pectinidens Dix. in Trans. R. Soc. S. Afr.
18: 251 (1930). Type: Transkei, Port St Johns, Wager
936 (BM, holo. ! ; PRE!).
Plants small, scattered or in small groups,
light green; terricolous. Stems 1-3 mm tall,
simple; in section round, central strand small,
inner cortical cells lax, outer cortical cells in
1-2 rows, smaller, weakly incrassate. Leaves
not crowded, little altered dry; lanceolate to
elliptical, 0,6-0, 8 mm long; apex acute,
cuspidate; margins irregular to serrulate,
elimbate or occasionally subperichaetial
leaves with a few differentiated cells on
vaginant laminae; costa percurrent. Vaginant
laminae to § of leaf length, termination of one
blade laminal; dorsal lamina gradually
tapering proximally to insertion, frequently
not reaching stem. Upper laminal cells irregu-
larly hexagonal to elongate-hexagonal, 12-17
pm long, smooth; basal cells of vaginant
laminae rectangular.
Perichaetia terminal; leaves curved,
asymmetrically broadly ovate-lingulate, 1,2
mm long, vaginant laminae limbate. Seta to 3
mm long, yellowish ; capsule erect, urn short-
cylindrical, 0,8 mm long, yellow-brown,
reddish at mouth; exothecial cells short-
rectangular to quadrate, longitudinal walls
thickened; peristome not seen; operculum
rostrate; spores round, 22-28 pm, weakly
papillose. Fig. 12: 22-28.
Map 19. — • Fissidens fasciculatus
x Fissidens parvilimbatus
Recently reported from Tanzania (Bizot & Poes,
1979). In Southern Africa the species has been col-
lected in the central Transvaal, Lesotho, Natal,
Transkei and the Cape. Map 19.
Voucher: Bosman 1611.
The presence of limbidia on the vaginant laminae of
perichaetial and subperichaetial leaves, large, smooth
leaf cells and rectangular basal leaf cells will identify
F. parvilimbatus. This species is similar to F. micro-
androgynus; see note under that species.
18. Fissidens pygmaeus Hornsch. in
Linnaea 15: 147 (1841); Broth, in Natiirl.
PflFam. 10: 148 (1924). Type: Cape, Table
Mountain, Ecklon s.n. (H-BR!).
Fissidens bifrons Schimp. ex C. Mull, in Bot. Ztg
17: 198 (1859); Broth, in Natiirl. PflFam. 10: 148
(1924); Sim, Bryo. S. Afr. 189 (1926). Type: Cape,
Green Kloof, Breutel s.n. (BM!).
Fissidens perpaucifolius Dix. ex Sim in Trans. R.
Soc. S. Afr. 15: 196 (1926). Type: Cape, Stellenbosch,
Wager 647 (BM, holo.!; PRE!).
Plants very small, scattered, yellow-green ;
terricolous. Stems to 4 mm high in sterile
plants, in fertile plants generally less than 1
mm high, simple; in section elliptical, central
strand present, cortical cells in 2-3 rows,
small, thickened. Leaves crowded in fertile
plants, distant in sterile plants, little altered
Fig. 12. — Fissidens submarginatus (1-8): 1. habit, xl; 2. habit, xlO; 3-4. leaves, X35; 5. cells at lower
margin of vaginant lamina, x435; 6. cells at lower margin of dorsal lamina, x435; 7. upper laminal cells at
margin, X640; 8. leaf apex, x435. F. scleromitrius (9—15) : 9. habit, xl; 10. habit, XlO; 11. stem in cross
section, x230; 12. leaf, x35; 13. cells at lower margin of vaginant lamina, x435; 14. cells at margin of dorsal
lamina, x640; 15. leaf apex, x435. F. microandrogynus (16-21): 16. habit, xl; 17. habit, XlO; 18. leaf, x35;
19. leaf base, X 170; 20. cells at margin of dorsal lamina, x640; 21. leaf apex, x 170. F. parvilimbatus (22-28): 22.
habit, xl; 23. habit, xlO; 24-25. leaves, x35; 26. leaf base, x435; 27. upper laminal cells, x640; 28. leaf
apex, X435. (1-8, Magill 5421 ; 9-15, Sim 9906; 16-21, Wager 79; 22-28, Sim 9905).
58
Fissidentaceae
dry; asymmetrically broadly ovate to ligulate,
0,3-0, 5 mm long; apex acute, generally bent
downwards in sterile plants; margins entire,
elimbate in lower leaves, limbate by 1-4 rows
of elongate cells on vaginant laminae of
subperichaetial leaves; costa strong, percur-
rent to very short-excurrent. Vaginant laminae
extending to upper leaf, termination of one
blade costal; dorsal lamina very narrow or
bulging distally in leaves of fertile plants,
ending well above insertion. Laminal cells
quadrate to short-rectangular, 10-15 pm long,
smooth, flat.
Perichaetia terminal; leaves asymmetri-
cal, broadly ovate, acuminate, to 1 mm long,
vaginant laminae distinctly limbate. Seta 3-7
mm long, yellowish; capsule erect to inclined,
urn short-cylindrical, 0,6-0, 8 mm long,
yellowish, mouth reddish; exothecial cells
quadrate, longitudinal walls thickened; peri-
stome deeply cleft, reddish; operculum short-
rostrate, 0,3 mm long; spores round, 22-25
pm , weakly papillose. Fig. 14: 1-9.
Endemic to Southern Africa, F. pygmaeus is
found in the northwestern and southwestern Cape
Province. Map 18.
Vouchers: Magill & Schelpe 3936, 3961 ; Rehmann
295 , 585; Schelpe 7731.
Sim (1926) was incorrect in his inclusion of F.
pygmaeus (=F. bifrons) in the synonomy of F.
splachnifolius. The latter is a very distinctive species,
belonging to the section Areofissidens, and has been
rarely collected. Fissidens pygmaeus is a more common
species of the section Semilimbidium.
Fissidens pygmaeus exhibits two distinct growth-
forms, i.e. elongate sterile plants with distant leaves
and short fertile plants with only a few broad,
crowded leaves. Most collections contain both sterile
and fertile plants.
Although direct contact between sterile and
fertile stems has not been observed, leaf shape,
anatomy and laminal cell size and pattern illustrate
the relationship between the two growth-forms. One
distinctive character of the sterile plants is the re-
flexed leaf apices. This character was also observed
on some of the small basal leaves of fertile plants,
although it was generally less pronounced.
19. Fissidens borgenii Hampe in Bot. Ztg
28: 36 (1870); Broth, in Naturl. PflFam. 10:
151 (1924). Type: Natal, Umpumulo, Borgen
s.n., 1867 (BM, holo.!; MANCH!).
Fissidens linearicaulis Broth. & Bryhn in Forh.
Vidensk Selsk. Krist. 1911 (4): 9 (1911); Broth, in
Naturl. PflFam. 10: 149 (1924). Type: Natal, Ekombe,
Titlestad s.n., Nov. 1907 (H-BR, holo.!).
Fissidens haakonii Broth. & Bryhn in Forh. Vidensk
Selsk. Krist. 1911 (4): 10 (1911); Broth, in Naturl.
PflFam. 10: 151 (1924). Type: Natal, Eshowe,
H. Bryhn s.n., Apr. 1908 (H-BR, holo.!).
Plants small, forming groups, yellow-
green to green; corticolous or rarely saxi-
colous. Stems 2-3 mm tall, simple; in section
subround, central strand present, inner cor-
tical cells large, in 2 rows, outer cortical cells
in 2-3 rows, stereids or substereids. Leaves
larger above, apically recurved dry, erect
wet; ligulate to asymmetrically oval-lingulate,
0,9-1, 4 mm long; apex acute to broadly acute;
margins minutely crenulate by marginal cell
papillae, elimbate or subperichaetial leaves
limbate on vaginant laminae only; costa
percurrent or ending just below apex. Vagi-
nant laminae to mid-leaf or just above,
termination of one blade submarginal to
laminal; dorsal lamina tapering proximally,
abruptly rounded at insertion. Laminal cells
hexagonal to quadrate, 5-10 pm long,
obscured by numerous, low, dense, peripheral
papillae.
Perichaetia terminal ; leaves 1 , 5 mm long,
limbidia on vaginant laminae only, laminal
cells elongate to rectangular, smooth. Seta
1-2 mm long, yellow; capsule erect to in-
clining, urn short-cylindrical, 0,5-0, 8 mm
long, strongly constricted below mouth when
dry, yellowish; exothecial cells quadrate,
longitudinal walls thickened ; peristome deep-
ly cleft, reddish; operculum rostrate; spores
round, 10-12 pm, weakly papillose. Fig. 14:
10-16.
Known from southeastern and Southern Africa,
F. borgenii is infrequently collected in dry woodlands
of the northern and eastern Transvaal, Zululand,
Natal, Transkei and the southwestern Cape. Map 15.
Vouchers: Magill 5222, 6164; Smook 1365;
Van Rooy 238 ; Von Breitenbach 220.
A distinctive species, F. borgenii is identified by its
acute apices, variable, weak limbidia restricted to the
vaginant laminae of perichaetial and subperichaetial
leaves, percurrent to short-excurrent costa and low,
dense papillae that obscure the leaf cells. Fissidens
borgenii is closely related to F. subobtusatus and F.
erosulus. In addition to the above characters, pre-
ference of rough-bark substrates will help to identify
F. borgenii.
20. Fissidens erosulus (C. Mull.) Par., Ind.
Bryol. 467 (1896); Broth, in Naturl. PflFam.
10: 151 (1924); Sim, Bryo. S. Afr. 191 (1926).
Type: Central Africa, Niam-Niam Region,
Nabambisso, Schweinfurth s.n., 17 Feb. 1870
(BM!).
Conomitrium erosulum C. Mull, in Linnaea 39: 367
(1875).
Fissidentaceae
59
Plants small, in groups, yellow-green to
dark green; terricolous. Stems (2-) 4-6 mm
tall, simple; in section round, central strand
present, inner cortical cells large, in single
row, outer cortical cells in 1 -2 rows, smaller,
incrassate. Leaves crisped or twisted dry, erect
wet; ligulate to Ungulate, somewhat asymme-
trical, 0,8-1 ,4 mm long; apex acute; margins
crenulate to serrulate by projecting cells,
elimbate or with weak limbidia on vaginant
laminae only, limbidia frequently consisting
of only a few elongated, smooth cells; costa
percurrent or ending just below apex. Vagi-
nant laminae extending to $ of leaf length,
termination of one blade laminal; dorsal
lamina gradually tapering to insertion. Lami-
nal cells hexagonal to quadrate, 5-7 n m long,
frequently bulging with 4-6 low, simple
papillae over lumen, not obscuring cells,
occasionally cells almost smooth.
Fig. 13. — Fissidens erosulus: 1. habit, X 1 ; 2.
habit, x 10; 3. leaf, x40; 4. leaf base, xl70; 5.
upper laminal cells (papillae partly shown), x640;
6. leaf apex, x 170; 7. capsule, x 15; 8. calyptra, x 15.
(1-8, Taylor 462).
Perichaetia terminal; leaves somewhat
asymmetrical, lanceolate, 1 , 5 mm long,
vaginant laminae limbate. Seta 2 , 5 mm long,
yellowish; capsule inclined, urn short-cylin-
drical, 0,5-0, 6 mm long; exothecial cells
quadrate, longitudinal walls thickened; peri-
stome reddish yellow, deeply cleft; operculum
rostrate; spores round, 10-12 n m, spiculate.
Fig. 13: 1-8.
Fissidens erosulus is known from central, eastern
and Southern Africa. In the Flora area it occurs in
northern Botswana, the northern, central and eastern
Transvaal, Swaziland and Zululand. Map 7.
Vouchers: Cholnoky 445; Magill 3592, 4984,
5366; Taylor 462; Van Rooy 216.
This species is very closely related to F. borgenii.
The two species can only be separated by leaf cell
ornamentation and substrate preferences. The con-
nection between substrate and papillae development
requires further study before the relationship between
these two species can be properly assessed.
The type specimen of F. erosulus has mammil-
lose upper lamina cells, but the basal leaf cells have
several low, papillae over each lumen. This corres-
ponds with several of the Zimbabwe specimens that
were examined. The cells of the Southern African
specimens are occasionally weakly bulging and have
4-6 low, simple papillae in a central ‘ring’ over each
lumen. Unlike the peripheral papillae of F. borgenii,
these papillae do not obscure the leaf cells.
21. Fissidens subobtusatus C. Mull, in
Hedwigia 38: 56 (1899); Broth, in Naturl.
PflFam. 10: 151 (1924). Type: Transvaal,
Lake Chrissie, Wilms s.n., Apr. 1885, in herb.
Jack (G. holo.!).
Fissidens borgenii Hampe var. obtusifolius Dix. in
Trans. R. Soc. S. Afr. 8: 187 (1920); Sim, Bryo. S.
Afr. 192 (1926). Type: Natal, Van Reenen, Wager 166
(PRE!).
Plants small, forming loose groups, light
green; terricolous. Stems 2-3 mm long,
simple; in section subround, central strand
present, inner cortical cells in 3 rows, small,
outer cortical cells in 2 rows, stereids or
substereids. Leaves larger above, contorted
dry, erect-spreading wet; ligulate to lingulate,
0,4-0, 7 mm long; apex broadly acute, obtuse
or rounded, abruptly apiculate by a smooth
clear cell; margins elimbate, crenulate; costa
ending 6-8 cells below apex. Vaginant laminae
extending to mid-leaf or just above, termin-
ation of one blade laminal; dorsal lamina
gradually tapering to insertion. Laminal cells
quadrate to hexagonal, 6-8 //m long, with 4-6
low, blunt papillae over lumen.
Perichaetia terminal; leaves ligulate, to
1 mm long, vaginant laminae limbate with a
60
Fissidentaceae
JfU
nyf jt
djT
r i \
■-nW
rtnJ,
i
111
Fissidentaceae
61
few elongate, smooth cells on margin. Seta
2, 0-2, 5 mm long, dark yellow; capsule
inclined, urn short-cylindrical, 0,8 mm long,
reddish yellow; exothecial cells quadrate,
longitudinal walls strongly thickened; peri-
stome deeply cleft, red-yellow; operculum
rostrate, beak ± oblique; calyptra cucullate;
spores round, 12-16 /tm, very weakly papil-
lose, green. Fig. 14: 17-23.
F. subobtusatus is found in grasslands of the
Transkei, northeastern Orange Free State, central
and eastern Transvaal and central South West
Africa/Namibia. It has also been reported from east
Africa (Bizot & Poes, 1979). Map 20.
Vouchers: Magill 3005; Scott 11135; Van Vuuren
1698.
Close to F. borgenii but differing in the smaller
size of the plants, shorter leaves with rounded apices
and the costa ending below the apex. In addition, F.
subobtusatus grows on soil and dolomite rock.
4. Section Crenularia
Crenularia C. Mm//, emend. Norkett ex Gangulee, Moss. E. India 456 (1971).
Section Crenularia C. Mull., Gen. Muse. Fr. 62 (1900).
Plants small; corticolous. Leaves without limbidia. Laminal cells small, papillose or
mammillose.
22. Fissidens pseudoserratus (C. Mm//.)
Jaeg., Enum. Fissid. 24 (1869); Broth, in
Naturl. PflFam. 10: 151 (1924); Sim, Bryo. S.
Afr. 190 (1926). Type: Cape, Krakakamma,
Ecklon s.n., July 1852 (BM!).
Conomitrium pseudoserratum C. Mull, in Bot. Ztg
17: 197 (1859).
Plants small, scattered or in loose patches,
light green; corticolous or rarely saxicolous.
Stems 1,0-1, 5 mm tall, simple; in section
subround, central strand very small, inner
cortical cells in 1-2 rows, large, outer cortical
cells in 1-2 rows, small, incrassate. Leaves
flattened, with recurved apices dry, erect-
spreading wet; lanceolate, 0,6-0, 9 mm long;
apex acute; margins elimbate, serrate, fre-
quently dentate on vaginant laminae; costa
strong, percurrent. Vaginant laminae extend-
ing to mid-leaf, termination of one blade
laminal; dorsal lamina tapering proximally,
just reaching insertion. Laminal cells hexa-
gonal to angular, 6-1 2^m long, thickened,
weakly bulging to sharply mammillose.
Perichaetia terminal; leaves undifferen-
tiated. Seta 2 mm long, yellow ; capsule erect,
urn short-cylindrical, 0,3-0, 5 mm long,
yellowish; exothecial cells quadrate, bulging,
Map 20. — • Fissidens pseudoserratus
x Fissidens subobtusatus
Fig. 14. — Fissidens pygmaeus (1-9): 1. habit, x 1 ; 2. fertile plant, x 10; 3. sterile plant, x 10; 4. leaf from
fertile plant, x 100; 5. leaf from sterile plant, x 100; 6. leaf base, x435; 7. upper laminal cells, x640; 8. leaf
apex, x435; 9. part of capsule mouth with peristome teeth, x 170. F. borgenii (10-16): 10. habit, X 1; 11. habit,
x 10; 12. leaf, x30; 13. leaf base, x435; 14. upper laminal cells (papillae partly shown), x640; 15. leaf apex,
x435; 16. capsule, x20. F. subobtusatus (17-23): 17. habit, xl; 18. habit, x 10; 19. leaf, x75; 20. cells at
base of vaginant laminae, x435; 21. cells at base of dorsal lamina, x435; 22. upper laminal cells (papillae
partly shown), x 640; 23. leaf apex, x435. F. pseudoserratus (24-30): 24. habit, x 1 ; 25. habit, x 10; 26. leaf,
xlOO; 27. leaf base, x435; 28. upper laminal cells at margin, x640; 29. leaf apex, x435; 30. capsule, x20,
(1, 3, 5, 9, Breutel s.n.; 2, 4 & 6-8, Ecklon s.n.; 10-16, Van Rooy 98; 17-23, Van Vuuren 1698; 24-25 & 30.
Schelpe 4407; 26-29, Sim PRE-CH5096).
62
Fissidentaceae
thin-walled; peristome deeply cleft, reddish;
spores round, 12-14 //m, weakly papillose.
Fig. 14: 24-30.
Endemic to Southern Africa, the species occurs
in grasslands and woodlands of the central Transvaal,
Zululand, Natal and eastern Cape. Map 20.
Vouchers: Crosby 7785; Magill 3125, 5571;
Schelpe 4407.
The small plants with narrow, elimbate leaves,
strongly serrate margins and mammillose leaf cells
will identify F. pseudoserratus. The species is quite
distinct and is the only representative of section
Crenularia in Southern Africa.
5. Section Aloma
Aloma C. Miill., Gen. Muse. Fr. 61 (1900).
Plants small; terricolous. Leaves without limbidia; costa percurrent to short-excurrent.
Laminal cells smooth to convex.
23. Fissidens nitens Rehm. ex Salm. in
Ann. Bot. 13: 121 (1899); Broth, in Natiirl.
PflFam. 10: 149 (1924); Sim, Bryo. S. Afr.
192 (1926). Type: Natal, Inanda, Rehmann
289 (BM!).
Plants small, in loose groups, yellow-
green, glossy; semi-aquatic. Stems 3-4 mm
tall, simple; in section elliptical, central strand
absent, inner cortical cells large, outer cortical
cells large, 2-3 rows, smaller, incrassate or sub-
stereids. Leaves equally spaced, little altered
dry, erect-spreading wet; broadly lanceolate
to ovate-lingulate, 1,0-1 ,2 mm long; apex
acute; margins bistratose; in section forming
4-celled, club-shaped border, minutely serru-
late above, marginal cells similar to laminal
cells; costa subpercurrent. Vaginant laminae
extending to mid-leaf, termination of one
blade laminal; dorsal lamina very gradually
tapering proximally to abruptly rounded at
base. Laminal cells hexagonal to subhexa-
gonal, 12-15 /urn long, incrassate, smooth;
basal cells undifferentiated.
Sporophyte not known. Fig. 1 1 : 24-30.
In Southern Africa, specimens of F. nitens have
been collected in or near running water in Natal and
Zululand. The species was recently reported from east
Africa by Bizot & Poes (1979). Map 17.
Voucher : Van der Plank 240.
The bistratose margins of this species are obvious
in specimens mounted in Hoyer’s medium. Water-
mounted specimens generally exhibit a darker margin.
Since the marginal cells of F. nitens are not
differentiated from the laminal cells, the bistratose
margin is not interpreted as a true limbidium. The
species is therefore placed into section Aloma.
6. Section Crispidum
Crispidum C. Miill. emend. Norkett ex Gangulee, Moss. E. India 457 (1971).
Section Crispidum C. Miill., Gen. Muse. Fr. 64 (1900).
Plants medium to large; terricolous. Stems with axillary hyaline nodules between vaginant
laminae. Leaves without limbidia, margins often crenulate. Laminal cells rounded, incrassate,
frequently mammillose.
24. Fissidens glaucescens Hornsch. in
Linnaea 15: 154 (1841); Broth, in Natiirl.
PflFam. 10: 151 (1924); Sim, Bryo. S. Afr.
201 (1926). Type: Cape, Table Mountain,
Ecklon s.n. (H-BR!).
Fissidens mucronatus Schimp. ex C. Miill. in Bot.
Ztg 16: 154 (1858). Type: Cape, Bethal, Breutel s.n.
(BM, holo.!).
Fissidens lanceolatus Hampe in Bot. Ztg 28: 36
(1870); Broth, in Natiirl. PflFam. 10: 151 (1924).
Type: Natal, Mapumulo, Borgen s.n., Jan. 1867
(BM, holo.!).
Fissidens cymatophyllus C. Miill. in Linnaea 42:
238 (1879); Bizot in Bull. Soc. bot. Fr. 114: 423(1967).
Type: Cape, Somerset East, Boschberg, MacOwan
s.n., 1878 (GRA!).
Fissidens rehmannii C. Miill. in Hedwigia 38: 56
(1899); Broth, in Natiirl. PflFam. 10: 151 (1924).
Type: Natal, Inanda, Rehmann 282d (NH!).
Fissidens procerior Broth. & Bryhn in Forh. Vidensk
Selsk. Krist. 1911: 9 (1911). Type: Natal, Entumeni,
H. Bryhn s.n., Apr. 1908 (H-BR!).
Fissidens zuluensis Broth. & Bryhn in Forh. Vidensk
Selsk. Krist. 1911: 9 (1911). Type: Natal, Entumeni,
H. Bryhn s.n., Apr. 1908 (H-BR!).
Fissidentaceae
63
Plants large, in loose groups, light green;
terricolous, frequently semi-aquatic. Stems
5-10 mm long, occasionally branched; in
section elliptical, central strand present, inner
cortical cells large, in 3 (4) rows, outer cortical
cells stereids, in 3 rows, reddish; axillary
hyaline nodules numerous, of 6-10 enlarged,
clear or chlorophyllous cells, enclosed by
vaginant laminae. Leaves equally spaced, erect
with circinate tips dry, erect-spreading wet,
occasionally somewhat undulate; oblong to
oblong-lanceolate, 2, 0-3, 2 mm long; apex
obtuse to broadly acute, cuspidate; margins
elimbate, entire to serrulate, frequently serrate
at apex; costa strong, short-excurrent, flex-
uous above. Vaginant laminae f of leaf length,
termination of one blade submarginal or
laminal; dorsal lamina not narrowing toward
stem, rounded proximally, generally crumpled
against stem. Upper laminal cells rounded-
hexagonal, 8-1 2 pm long, incrassate, mammil-
lose; quadrate marginally, smooth; cells of
vaginant laminae quadrate to rectangular,
smooth, becoming smaller toward margin.
Dioicous. Perigonia axillary, gemmate.
Perichaetia terminal or infrequently axillary;
leaves not differentiated. Seta to 10 mm long,
reddish yellow; capsule horizontal, urn
curved, 1,2-1, 5 mm long, red-yellow; exo-
thecial cells rectangular, incrassate ; peristome
teeth deeply cleft, red-yellow; operculum
conic-rostrate, 0,6 mm high; spores round,
15-20 pm, smooth. Fig. 15: 1-6.
The species is known from eastern and Southern
Africa. It is the most widely distributed species of
Fissidens in the Flora area and occurs in grassland
Map 21. — • Fissidens glaucescens
and woodland of the northern, eastern, central and
southern Transvaal, Swaziland, Zululand, Natal,
Lesotho, Orange Free State, Transkei and the eastern,
central, southern and western Cape. Map 21.
Vouchers: Cholnoky 410; Esterhuysen 24, 444;
Magill 3085, 4763, 5622; Schelpe 2033; Van Rooy 84.
The large size of the plants, leaf shape and small,
incrassate, mammillose leaf cells will separate speci-
mens of F. glaucescens. The species is similar to F.
asplenioides , but the two species are easily divided
by the shape of their leaf apices, costal development
and morphology of the dorsal laminae. Intermediates
between the two species were not seen.
F. glaucescens is included in section Crispidium
because of the axillary hyaline nodules regularly
produced on the stem between the vaginant laminae.
Occasionally meristematic activity was observed in
these structures, resulting in the development of peri-
chaetial or perigonial buds; vide Iwatsuki & Pursell,
1980.
7. Section Serridium
Serridium C. Mull, emend. Norkett ex Gangulee, Moss. E. India 457 (1971).
Section Serridium C. Mull., Gen. Muse. Fr. 67 (1900).
Plants large to robust; terricolous. Stems without axillary hyaline nodules. Leaves' without
limbidia, marginal cells occasionally weakly differentiated. Laminal cells rounded, incrassate,
generally mammillose.
25. Fissidens plumosus Hornsch. in Lin-
naea 15: 151 (1841); Broth, in Natiirl. PflFam.
10: 151 (1924); Sim, Bryo. S. Afr. 203 (1926).
Type: Cape, Vorgebirge, Mundt & Maire s.n.
Plants large, in small groups; terricolous
or saxicolous. Stems 5-7 mm long, un-
branched; in section elliptical, central strand
present, inner cortical cells in 2-3 rows, large,
incrassate, outer cells in 2-3 rows, stereids.
Leaves crowded, recurved to secund wet or
dry; linear, (2-)3-5 mm long; apex acute;
margins elimbate, smooth below, crenulate to
serrate above ; costa percurrent to mucronate.
Vaginant laminae | of leaf length, termination
64
Fissidentaceae
Fissidentaceae
65
of one blade marginal to submarginal ; dorsal
lamina tapering proximally, generally not
reaching stem, if so, decurrent by single row
of cells. Upper laminal cells rounded, mixture
of large (12-20 pm long) and small (7-10 pm
long) cells, quadrate to short-rectangular,
strongly incrassate, weakly mammillose to
smooth; cells of vaginant laminae similar but
becoming smaller toward margin.
Dioicous. Perichaetia terminal; leaves
undifferentiated. Seta 8-9 mm long, yellow-
brown, bent upward from prostrate stem;
capsule horizontal, urn oval, 1 mm long,
reddish yellow; exothecial cells rectangular,
incrassate; peristome teeth cleft to middle,
reddish yellow; operculum conic-rostrate, to
0,5 mm long; spores round, 12-13 pm,
spiculate. Fig. 15: 7-12.
Endemic to Southern Africa, F. plumosus is
frequently collected on shaded banks in forests and
fynbos of the southern and southwestern Cape. A
few collections have also been made in the north-
western and eastern Cape, Natal and the central,
eastern and northern Transvaal. Map 22.
Vouchers: Esterhuysen 26903; Magill 6192;
Smook & Phelan 839; Schelpe 7856.
The habit, narrow leaves and large, flat leaf
cells should identify this species. The development of
the dorsal lamina is useful in separating the species
from F. glaucescens.
Some confusion between F. plumosus and F.
fasciculatus has been noted on earlier collections. The
habit and leaf shape of the two species are quite
similar, but F. plumosus is a much shorter plant and
its leaves are unistratose throughout.
26. Fissidens asplenioides Hedw., Spec.
Muse. 156 (1801); Broth, in Nattirl. PflFam.
10: 151 (1924); Grout in N. Amer. FI. 15, 3:
192(1943). Type: Jamaica, s. coll. (G, holo.!).
Fissidens amblyophyllus C. Mull, in Hedwigia 38:
57 (1899); Broth, in Natiirl. PflFam. 10: 151 (1924);
Sim, Bryo. S. Afr. 203 (1926). Syntypes: Natal, Van
Reenens Pass, Rehmann 285b (PRE!); Inanda,
Rehmann 285 (PRE!); Transvaal, Spitzkop, Wilms
s.n., 1887 (G!).
Plants large, forming turfs or groups,
yellow-green to light green; terricolous. Stems
10-15 mm tall, infrequently branched; in
section subround, central strand small, inner
cortical cells large, in 3 rows smaller toward
Map 22. — • Fissidens plumosus
X Nanobryum dummeri
margin, outer cortical cells in 2 rows, stereids,
red-yellow. Leaves somewhat crowded, erect
with circinate apices dry, patent wet ; ligulate
to oblong-lanceolate, 2-3 mm long; apex
obtuse to rounded, rarely broadly acute;
margins elimbate, crenulate; costa ending
below apex, flexuose above. Vaginant laminae
to j leaf length, one blade abruptly rounded
distally, terminating on lamina or costa;
dorsal lamina abruptly rounded near inser-
tion. Laminal cells rounded, hexagonal to
subhexagonal, 8-10 pm long, incrassate,
superficially bulging; slightly larger in vagi-
nant laminae.
Perichaetia terminal or lateral; leaves
undifferentiated. Seta 3-5 mm long; capsule
erect, urn short-cylindrical, 0, 5-1,0 mm
long; peristome deeply cleft, reddish; oper-
culum long-rostrate; spores not seen. Fig. 15:
13-18.
Fissidens asplenioides is widely distributed,
especially in the Southern Hemisphere. The species is
known from Central and South America, Africa,
India, southeast Asia, Australia and New Zealand.
In Southern Africa, the species is found in grassland
and woodland communities of the northern, central
and eastern Transvaal, Zululand, Natal, northeastern
Orange Free State as well as the eastern and southern
Cape. Map 10.
Fig. 15. — Fissidens glaucescens (1-6): 1. habit, xl;2. habit, x6;3. leaf, x50;4. leaf base, xl70;5. leaf
apex, xl70; 6. upper laminal cells, x640. F. plumosus (7-12): 7. habit, X 1 ; 8. habit, x6; 9. leaf, x50; 10.
leaf base, xl70; 11. leaf apex, xl70; 12. upper laminal cells, x640. F. asplenioides (13—18) : 13. habit, xl;
14. habit, x6; 15. leaf, x50; 16. leaf base, xl70; 17. leaf apex, xl70; 18. upper laminal cells, x640. (1-6,
Cholnoky 410; 7-8, Schelpe 7856; 9-12, Magill 6032; 13-18, Magill 3054).
66
Fissidentaceae
4
Fissidentaceae
67
Vouchers: Cholnoky 183, 348; Kluge 1022;
Rankin 77 ; Schelpe 4406; Van Rooy 165.
The obtuse to rounded leaf apices, costa ending
below apex and dorsal laminae ending above the
insertion distinguish F. asplenioides in Southern
Africa. The large plants and incrassate, mammillose
leaf cells suggest a close relationship to F. glaucescens ;
see note under that species.
3. Subgenus Pachyfissidens
Pachyfissidens (C. Mull.) Kindb., Eur. N. Amer. Bryin. 2; 165 (1897).
Section Pachyfissidens C. Mull., Syn. Muse. 1 : 45 (1848).
Plants aquatic or semi-aquatic. Stems without central strand. Leaves costate, laminae
bistratose.
27. Fissidens fasciculatus Hornsch. in
Linnaea 15: 155 (1841); Broth, in Nattirl.
PflFam. 10: 153 (1924); Sim, Bryo. S. Afr.
202 (1926). Type: Cape, Du Toit’s Kloof,
Schonfeld, Drege s.n., 1828 (BM!).
Plants large, forming patches, dark green
to blackish green; terricolous or saxicolous,
aquatic to semi-aquatic. Stems 10-20 mm
long, unbranched ; in section elliptical, central
strand present, inner cortical cells large,
incrassate, in 3-4 rows, outer cortical cells
smaller, strongly incrassate, marginal row
stereids or substereids. Leaves crowded, larger
above, appressed, curved dry, appressed to
erect wet, linear-lanceolate, 3-5 mm long,
bistratose above and in dorsal laminae ; apex
long-acuminate; margins elimbate, plane,
entire; costa percurrent or ending just below
apex. Vaginant laminae of leaf length,
unistratose, termination of one blade laminal ;
dorsal lamina bistratose, tapering proximally,
generally ending well above insertion. Lami-
nal cells rounded, quadrate to rectangular or
angular, 10-17 /tm long, incrassate, smooth.
Dioicous. Perichaetia terminal; leaves
undifferentiated. Seta 5-6 mm long, yellowish;
capsule erect, urn short-cylindrical, to 1 mm
long, red-yellow; exothecial cells quadrate to
rectangular, thickened; peristome teeth deeply
cleft, reddish; operculum rostrate; spores
(immature) round, 12-15 //m, smooth. Fig.
16: 1-7.
Endemic to Southern Africa. F. fasciculatus is
found in or around streams in the southern and south-
western Cape. Map 19.
Vouchers: Cholnoky 360; Crosby & Crosby
9257 ; Esterhuysen 21229; Magill 6213.
The large, blackish green plants, narrow leaves
and bistratose upper leaf and dorsal lamina characte-
rize F. fasciculatus. The plants are somewhat similar
to F. plumosus but the two species are easily separated,
as the leaves of the latter are broader and unistratose
throughout ; see note under that species.
4. Subgenus Octodiceras
Octodiceras (Brid.) Broth, in Naturl. PflFam. 1: 361 (1901).
Genus Octodiceras Brid., Muscol. Recent. Suppl. 1: 162 (1806).
Plants aquatic, floating. Stems without central strand. Leaves costate; laminae unistratose.
28. Fissidens fontanus ( B . Pyl.) Steud.,
Nom. Bot. 2: 166 (1824); Scott & Stone,
Moss. S. Aust. 86 (1976). Type: Europe.
Skitophyllum fontanum B. Pyl. in J. de Bot., ser 2,
4: 133 (1815). Octodiceras fontanum (B. Pyl.) Lindb.
in Ofvers. K. VetenskAkad. Forh. 20: 405 (1863);
Smith, Moss FI. Brit. Irel. 204 (1978).
Fontinalis juliana (Savi) Savi ex Cand., FI. Franc,
ed. 3, 6: 236 (1815). Octodiceras julianum (Cand.)
Brid., Bryol. Univ. 2: 678 (1827). Conomitrium
julianum (Cand.) Mont, in Annls Sci. nat. Bot., ser. 2,
8: 246 (1837); Sim, Bryo. S. Afr. 205 (1926). Fissidens
julianus (Cand.) Schimp. in Flora, Jena 21 : 271
(1838); Broth, in Naturl. PflFam. 10: 153 (1924);
Grout, Moss FI. N. Amer. 1: 23 (1936). Type:
Europe.
Fig. 16. — Fissidens fasciculatus (1-7): 1. habit, xl;2. habit, x5;3.1eaf, x 50; 4. leaf base, xl70;5.upper
laminal cells, x640; 6. cells at termination of vaginant lamina, x 170; 7. leaf apex, x 170. F. fontanus (8-13):
8. habit. xl;9. habit, x5; 10. leaf, x50; 11. leaf base, xl70; 12. upper laminal cells, x435; 13. leaf apex,
X 170. (1-2, Esterhuysen 21229; 3-7, Cholnoky 359; 8-10, Cholnoky 604; 11-13, Cholnoky 641).
68
Fissidentaceae
Conomitrium capensis C. Miill., Syn. Muse. 2:
524 (1851). Octodiceras capensis (C. Miill.) Jaeg. in
Verh. St. Gall, naturw. Ges. 1874-75: 135 (1876).
Fissidens canensis (C. Miill.) Broth, in Natiirl. PflFam.
10: 154 (1924). Type: Cape, Drege sub 9379a & b
(H-BR!).
Plants large to robust, flaccid, green,
yellow-green or blackish; aquatic, saxicolous.
Stems 50-100 mm long; in section subround,
central strand absent, inner cortical cells large,
irregularly thickened at corners, outer cortical
cells small, round, incrassate. Leaves distant,
spreading wet or dry; linear-lanceolate,
(3— )4— 7 mm long, somewhat flexuous above;
apex acute to obtuse; margins elimbate, plane,
entire to wavy; costa ending well below apex.
Vaginant laminae 3—3 of leaf length, equal or
termination of one blade submarginal; dorsal
lamina abruptly tapering proximally, not
reaching stem. Laminal cells irregular, rectan-
gular to hexagonal, 18-37 /urn long, 12-15 /urn
wide, thin-walled, flat; juxtacostal cells
somewhat larger, rectangular.
Autoicous, gametangia on short lateral
branches. Seta very short, 0,4 mm long;
capsule erect, urn elliptical, 0,5 mm long;
peristome cleft and perforated; operculum
rostrate; spores round, 18-20 /urn. Fig. 16:
8-13.
Fissidens fontanus grows in water and is known
from North, Central and southern South America,
Europe and Africa. In Southern Africa the species is
most common in Natal and Transkei, but specimens
have been collected in the eastern Orange Free State,
central and eastern Transvaal, Zululand, and the
northern, southern and southwestern Cape. Map 8.
Vouchers: Cholnoky 593; Esterhuysen 17956;
Leighton 3367.
The large, flaccid plants with long, distant,
unistratose leaves characterize F. fontanus. Consider-
ing the size and habit of these plants and their
aquatic nature, it is unlikely that this species could
be confused with any other species of Fissidens.
Fontinalis antipyretica bears some resemblance, but
the two species are easily separated on generic
characters.
Insufficiently Known Species
Fissidens longulus C. Miill. in Hedwigia 38: 56
(1899). Type: Cape, Somerset East, Boschberg,
MacOwan s.n., 1883. The type has not been seen,
however the description indicates a close resemblance
to F. pygmaeus Hornsch.
Fissidens macleanus Shaw in Cape Monthly Mag.
17: 314 (1878). Type: Cape, Graaff-Reinet, McLea
s.n. The Sim correspondence (PRE) indicates that the
Shaw collection was destroyed sometime after his
death. Attempts to locate the specimens in Southern
Africa have been unsuccessful and duplicates were
not found in the European herbaria likely to have
Shaw specimens. Sim (1926) considered the species
to be synonymous with F. curvatus Hornsch. (as F.
pycnophyllus), although he did not see the type.
Fissidens pauperrimus C. Mull, in Hedwigia 38:
54 (1899), horn, illeg., non C. Miill. (1881). Fissidens
rutenbergii Par., Ind. Bryol. Suppl. 164 (1900). Type:
Southern Africa, Rutenberg s.n., 1877. The type
specimen has not been seen. Some doubt exists on
the occurrence of this species in the Flora area. The
collector is known for his collections on Madagascar
and other East African Islands, but he is not cited as a
collector from Southern Africa.
69
NANOBRYACEAE
Plants small, scattered along a persistent protonema, yellow-green; terricolous or corti-
colous. Stems erect, central strand present. Leaves larger above, symmetrical; elliptical,
abruptly subulate; base clasping stem; margins erect, entire or with large blunt tooth at
shoulders. Costa present or absent, weak below, occasionally extending into subula. Upper
laminal cells rhomboidal, thin-walled; basal cells scarcely differentiated.
Dioicous. Male plants smaller. Perichaetia terminal. Seta cygneous; capsule small,
nodding or pendent, weakly asymmetric; peristome single, teeth 16, deeply cleft, red-yellow;
operculum long-rostrate; calyptra cucullate, covering beak of operculum only; spores small,
granulose.
Nanobryaceae is a monotypic family, known only from Africa.
NANOBRYUM
Nanobryum Dix. in J. Bot., Lond. 60: 101 (1922); Broth, in Natiirl. PflFam. 11: 525 (1925);
Sim, Bryo. S. Afr. 150 (1926); Schultze-Motel in Willdenowia 5: 386 (1969). Type species:
N. dummeri Dix.
With characters of the family.
The genus Nanobryum consists of a single species reported from moist forests in several areas of Africa,
south of the Sahara. In recent years the concept of the genus has apparently been altered from Dixon’s (1922)
description of plants intermediate between Fissidentaceae and Dicranaceae. It appears that several authors
have placed small Fissidens species under Nanobryum. Examination of syntypes of N. dummeri, from Southern
Africa, clearly supports Dixon’s original description.
Nanobryum dummeri Dix. in J. Bot.,
Lond. 60: 101 (1922); Broth, in Natiirl.
PflFam. 11: 525 (1925); Sim, Bryo. S. Afr.
150 (1926). Syntypes: Uganda, Mulange,
Dummer 4080a (BM); Kipayo, Dummer 1214;
Transkei, Port St Johns, Wager 955 (BM!;
PRE!).
Plants small, scattered, yellow-green;
terricolous. Protonema persistent. Stems 0,1-
0,2 mm tall, simple; in section round, central
strand weak, inner cortical cells large, in single
row, thin-walled, outer cortical cells smaller,
in 2 rows, incrassate, yellow-brown. Leaves
larger above, weakly flexuose dry, erect-
spreading wet, elliptical, 0, 4-1,0 mm long,
upper and perichaetial leaves abruptly subu-
late, shoulders frequently with large, blunt
tooth; base clasping stem; margins erect,
entire. Costa weak below, stronger above,
frequently extending into subula or complete-
ly absent in some leaves; in section round,
guide cells 2-4, large, incrassate, ventral cells
in single row, large, incrassate, dorsal cells in
2-3 rows, smaller, incrassate. Upper laminal
cells fusiform to rhomboidal, thin-walled,
smooth; basal cells rectangular, slightly
larger.
Dioicous. Perichaetia terminal; leaves
not differentiated. Seta cygneous, 2-3 mm
long, yellowish; capsule nodding or pendent,
asymmetrically ovoid, 0,5 mm long, yel-
lowish, mouth reddish; exothecial cells
quadrate, thickened at corners, smaller at
mouth; peristome well developed, teeth 16,
lanceolate, 0,3 mm long, cleft to near middle,
finely papillose between trabeculae below,
red-yellow, apical filaments spirally striate,
hyaline; operculum rostrate, 0,4-0, 6 mm
long; calyptra cucullate, 0,4 mm long,
covering operculum beak; spores round, 10-
12 pm, granulate, yellowish. Fig. 17: 1-12.
The species is infrequently reported from moist
forests of western, central, eastern and Southern
Africa. In the Flora area the species has been col-
lected only at Port St Johns in the Transkei. Map 22.
Voucher: Type only.
As suggested by Dixon, when he described the
species, N. dummeri is intermediate between Dicra-
nella and Fissidens. The sporophytes are practically
70
Nanobryaceae
identical to the sporophytes of Fissidens, including
peristome ornamentation. Gametophytically the
plants are more similar to Dicranella. The plants
examined showed a very symmetrical development
of the lamina and no indication of a dorsal lamina as
seen in Fissidens.
The plants from Oubangui, discussed and illus-
trated by R. Potier de la Varde (1927) under this
name, undoubtedly belong to a species of Fissidens
rather than to Nanobryum. Because of the apparent
confusion of the identity of N. dummeri in the northern
parts of its range, the combination N. gladiolum
(Mitt.) Bizot, has not been adopted for the Southern
African plants. It also appears from the illustration
that Mitten’s (1860) plant is properly placed in
Fissidens-, vide Bizot & Poes (19791.
Fig. 17. — Nanobryum dummeri: 1. habit, xl;
2. habit, x 10; 3. habit, x 20; 4. stem in cross section,
X 270; 5-7. leaves, x50; 8. leaf in proximal section,
x435; 9. leaf in distal section, x435; 10. cells at
mid-leaf, xl70; 11. upper leaf, Xl70; 12. peristome
teeth, x 170. (1-12, Wager 53).
71
ARCHIDIACEAE
by J. van Rooy
Plants small, yellow to yellow-green; terricolous. Stems erect or prostrate, often stoloni-
ferous, frequently branching by subperichaetial innovations; in section round, central strand
present, inner cortical cells large, outer cortical cells usually smaller. Leaves frequently reduced
to bracts below, primary stem and innovation leaves usually similar, but innovation leaves
smaller.
Autoicous. Perigonia gemmate, terminal on short lateral branches or axillary on stem.
Perichaetia gemmate, terminal or lateral; leaves erect to erect-spreading, frequently concave,
elliptic or oblong to ovate or lanceolate, apex apiculate, acute, acuminate or subulate; margins
plane to recurved, entire, serrulate, irregularly crenulate or papillose-denticulate. Costa in
section circular, semi-circular or crescent-shaped, cells undifferentiated or dorsal cells smaller,
incrassate or stereids. Laminal cells uniform or irregular in shape; basal cells rectangular,
thin-walled, larger, hyaline. Capsules subsessile or sessile, cleistocarpic; globose, wall single-
layered, undifferentiated, semi-transparent, yellowish, with brownish spots when mature,
stomata absent; calyptra undeveloped, inconspicuous; spores large, >50 pm, rounded to
polyhedral, thick-walled, smooth to densely papillose, pale yellow to brown, 4-180 per capsule.
During compilation of Archidiaceae, a revision of the genus Archidium (Snider, 1975), was consulted
freely and was invaluable in completing this study.
ARCHIDIUM
Archidium Brid., Bryol. Univ. 1: 747 (1827); Broth, in Natiirl. PflFam. 10: 155 (1924); Sim,
Bryo. S. Afr. 140 (1926); Snider in J. Hattori bot. Lab. 39: 122 (1975). Type species: A.
phascoides Brid.
With characters of the family.
Small plants forming cushions, turfs, or tufts on sandy or gravelly soils. The small size and yellowish
colour make Archidium difficult to see in the field. Many of the Southern African species are only represented
by a few specimens or even single collections. Sterile plants of Archidium can easily be mistaken for small
species of Bryum and therefore fruiting plants are generally needed for identification.
Archidium is cleistocarpic and the sessile or subsessile, globose capsule is surrounded by large perichaetial
leaves. All Southern African species of Archidium are autoicous and the capsules are terminal, except in A.
ohioense where the capsules are terminal or lateral. Only a few spores are produced in each capsule but they are
extremely large.
The greatest number of described species is found in Africa, where 14 of the c. 26 species of Archidium
occur. Eleven species occur in Southern Africa, seven are endemic to the Flora region. Of the endemic taxa
five are very restricted in distribution, and are presently known only from the southwestern Cape.
1 Spores < 100 /im in diameter, smooth, 75-178 spores per capsule (Subgenus Archidiella) 1. A. dinteri
1 Spores > 100 //m in diameter, smooth to densely papillose; 4—70 spores per capsule (Subgenus Archidium ):
2 Spores densely papillose 6. A. muelleranum
2 Spores smooth to granulose:
3 Most leaf margins irregularly denticulate; branch leaves appressed 9. A. julicaule
3 Leaf margins entire to crenulate; branch leaves generally erect-spreading:
4 Perichaetial leaf cells 40-100 //m long; costa in section circular:
5 Costa percurrent to short-excurrent :
6 Perichaetial leaf cells irregularly rectangular 5. A. subulatum
6 Perichaetial leaf cells fusiform to rhomboidal:
7 Upper stem leaf margins plane; leaves narrow; laminal cells regular in shape. . .. 2. A. ohioense
7 Upper stem leaf margins narrowly recurved; leaves broad; laminal cells irregular in shape
11. A. andersonianum
72
Archidiaceae
5 Costa excurrent, frequently forming distinct awn:
8 Laminal cells of upper stem leaves short-rhombic to rhomboidal; apex short-acuminate;
costa long excurrent 4. A. acanthophyllum
8 Laminal cells of upper stem leaves elongate, rhomboidal to fusiform; apex acuminate to long-
acuminate:
9 Costa long-excurrent; upper perichaetial leaf cells narrow, cell walls 3-7 //m wide
3 .A. microthecium
9 Costa excurrent; upper perichaetial leaf cells wide, cell walls <4 pm thick 2. A. ohioense
4 Perichaetial leaf cells 15 to 45 pm long, occasionally some cells longer; costa in section crescent-
shaped or semi-circular:
10 Costa excurrent as a distinct awn 8. A. capense
10 Costa percurrent:
11 Perichaetial leaves bistratose juxtacostally 7. A. rehmannii
1 1 Perichaetial leaves unistratose juxtacostally 10. A. amplexicaule
1. Archidium dinteri ( Irmsch .) Snider in
J. Hattori bot. Lab. 39: 126 (1975). Type:
South West Africa/Namibia, near Warmbad,
Dinter s.n., 1924 (HBG, holo.).
Archidiella dinteri Irmsch. in Mitt. Inst. allg.
Bot., Hamb. 6: 338 (1926).
Plants forming loose mats, yellow-green;
terricolous. Stems to 10 mm long, prostrate
or suberect, stoloniferous, branching; in
section central strand present, cortical cells
large, thin-walled, outer 1-2 rows smaller.
Leaves bract-like, appressed; short-acuminate
to apiculate, 0,2-0, 4 mm long; margins
plane, entire to denticulate. Costa subper-
current to percurrent, weak. Laminal cells
rhomboidal or rectangular, thin-walled.
Perichaetia terminal on erect branches.
Leaves erect, concave, ovate to obovate,
apex short-acuminate or apiculate, 0, 5-1,0
mm long; margins plane, entire or denticulate
above. Costa percurrent, weak; in section
semi-circular, cells undifferentiated, in 2
rows. Laminal cells irregularly rhomboidal
or rectangular, 30-50 pm long, thin-walled;
apical cells longer; basal marginal cells
hyaline. Spores 75-178 per capsule, irregularly
rounded to polyhedral, 48-82 pm, smooth,
yellow. Fig. 18: 1-10.
Map 23. — • Archidium dinteri
x Archidium mueileranum
Archidium dinteri is known from only two speci-
mens, collected at Warmbad and Dassiskuppe in
South West Africa/Namibia, on quartz sand. Map
23.
Voucher: Volk 6903.
The subgenus Archidiella (Irmsch.) Snider is
based on A. dinteri and can be separated from the
subgenus Archidium by the differences in spore size
and number.
Fig. 18. — Archidium dinteri (1-10): 1. habit, x 1 ; 2. habit showing capsule and spores, x40; 3. stem in
cross section, x435; 4. leaves, x40; 5. perichaetial leaves, x40; 6. perichaetial leaf in cross section, x435;
7. perichaetial leaf cells (upper left margin), xl70; 8. upper laminal cells of perichaetial leaf, x640; 9. peri-
chaetial leaf apex, xl70; 10. spores, xl70. A. ohioense (11-16): 11. habit, xl; 12. habit, x40; 13. leaves,
x 10; 14. perichaetial leaf, x40; 15. upper perichaetial leaf cells, x435; 16. cells in upper perichaetial leaf,
X 170. A. microthecium (17-22): 17. habit, xl; 18. habit, x40; 19. leaves, x40; 20. perichaetial leaf, x40;
21. upper perichaetial leaf cells, X435; 22. cells in upper perichaetial leaf, xl70. (1-6 & 8-10, Volk 6903; 7,
Dinter 1924; 11-12, Sim 10004; 13-16 Ma^V/3563; 17-22, Magill 4951).
Archidiaceae
74
Archidiaceae
2. Archidium ohioense Schimp. ex C.
Miill., Syn. Muse. 2: 517 (1851); Snider in J.
Hattori bot. Lab. 39: 135 (1975). Type:
United States of America, Sullivant, Musci
Allegh. 213 (FH, lecto.), vide Snider (1975).
Archidium africanum Mitt, in J. Linn. Soc., Bot.
22: 299 (1885); Sim, Bryo. S. Afr. 140 (1926). Type:
Tanzania, Usagara Mtns, Hanning ton s.n. (NY,
holo.).
Archidium pellucidum Dix. ex Sim, Bryo. S. Afr.
142 (1926). Type: South West Africa/Namibia,
Erongo Mtns, Pearson 9849 (BM, holo.; PRE!).
Archidium leptophyllum P. Varde in Revue bryol.
lichen. 59: 86 (1932). Type: Central African Republic,
Tisserant 566d (PC, holo.).
Plants forming loose turfs, yellow-green;
terricolous. Stems erect or subflexuose, 2-15
mm high, simple or branching; in section
with central strand, cortical cells large, thin-
walled, outer 1-2 rows smaller. Leaves
bract-like below, erect-spreading or wide-
spreading above, ovate-lanceolate to ovate or
occasionally narrowly lanceolate, short-acu-
minate to subulate, 0,9-1, 5 mm long; base
often clasping; margins plane, entire or
serrulate above. Costa percurrent to excurrent
as hair-point; in section cells undifferentiated.
Laminal cells rhomboidal to fusiform, basal
cells rectangular to quadrate.
Perichaetia terminal or lateral. Leaves
erect, ovate, oblong-ovate, ovate-lanceolate
or lanceolate, acuminate, 0,7-1, 8 mm long;
margins plane to recurved; entire to serrulate
above. Costa percurrent to excurrent as hair-
point; in section round, cells undifferentiated,
incrassate. Laminal cells rhomboidal to
fusiform, frequently lax, 37-95 pm long,
walls (1,7-) 2,8 ( — 4,4) pm wide, lumen
(7 — ) 12 ( — 15) pm wide; basal cells rhom-
boidal to rectangular, lax, hyaline below.
Spores 8-60 per capsule, rounded triangular
to polyhedral, 97-176 //m, smooth to granul-
ose, yellow. Fig. 18: 11-16.
This species is widely distributed and has been
collected in North America, West Indies, India, Sri
Lanka, Japan and Africa. In Southern Africa it is
known from the southwestern and southern Cape,
Zululand, Swaziland and the eastern and central
Transvaal and South West Africa/Namibia. The
plants grow on sandy soil in moist or dry habitats.
Map 24.
Vouchers: Magill 3563; Sim 10004; Wager 63.
Two growth forms of A. ohioense can be recog-
nized in Southern Africa. In the short ‘form’, branch-
ing is mostly by subperichaetial innovations; the stem
leaves are rapidly reduced to ovate-lanceolate bracts
below, and the perichaetial leaves are large, broadly
ovate-lanceolate and have percurrent to excurrent
costae. The tall, little branched, subflexuose ‘form’
has lateral perichaetia. The narrowly long-triangular
stem leaves are distant, widespreading and reduced
only near the stem base. The perichaetial leaves in
this ‘form’ are smaller with a more or less percurrent
costa. Although there is a complete gradient from the
short ‘form’ to the tall ‘form’ in North America
(Snider, 1975), this is not evident in Southern Africa.
The tall, subflexuose ‘form’ occurs in moist, shady
habitats and is known in Southern Africa from a
single specimen collected in the Ngoya Forest of
Zululand. The short ‘form’ is more common and
occurs in drier, open habitats.
3. Archidium microthecium Dix. & P.
Varde in Ann. Cryptog. Exot. 1: 37 (1928);
Snider in J. Hattori bot. Lab. 39: 141 (1975).
Type: India, Kodiakanal, Foreau 211 (BM,
holo.).
Plants caespitose, yellow or yellow-green;
terricolous. Stems 1-5 mm high, branching by
subperichaetial innovations; in section with
central strand of collapsed cells, inner cortical
cells thin-walled, large, in single row, outer
cortical cells smaller, in 1-2 rows. Leaves
bract-like below, larger above, erect to erect-
spreading, ovate-lanceolate to lanceolate,
acuminate, 0,9-1 , 1 mm long; margins plane,
entire. Costa excurrent or forming hair-point;
in section round, cells undifferentiated,
incrassate or stereids. Laminal cells rhom-
boidal to fusiform, incrassate; basal cells
short-rectangular to quadrate; lower marginal
cells rectangular to quadrate.
Archidiaceae
75
Perichaetia terminal. Leaves erect to
erect-spreading, occasionally secund; ovate-
lanceolate to oblong-lanceolate, acuminate,
0,8-1, 4 mm long; margins entire, plane to
faintly recurved above. Costa excurrent as
hair-point; in section round, surface cells
incrassate or substereids, central cells sub-
stereids to stereids. Laminal cells fusiform,
shorter above and along margins, 48-88 pm
long, walls 3-7 pm wide, lumen 5-9 pm
wide; basal cells rectangular, thin-walled,
basal marginal cells hyaline. Spores 10-17 per
capsule, irregularly rounded, 88-202 pm,
smooth to granulose, yellow. Fig. 18: 17-22.
This species has been collected on sandy soils in
South West Africa/Namibia, Natal and the Kruger
National Park in the northeastern Transvaal. The
plant also occurs in east Africa and southern and
eastern India. Map 25.
Map 25. — • Archidium microthecium
X Archidium subulatum
Vouchers: Magill 4948, 4997; Volk 01160.
Archidium microthecium is separated from the
closely related A. ohioense on the basis of its nar-
rower and more incrassate lamina cells, long-excur-
rent costa, and oblong-lanceolate perichaetial leaves.
4. Archidium acanthophyllum Snider in
Bryologist 78: 152 (1975). Type: Transvaal,
Kruger National Park, Pretoriuskop, Godfrey
GH. 1649b (DUKE, holo.).
Plants forming turfs, yellow to yellow-
green; terricolous. Stems erect or prostrate,
3-8 mm tall, branching; in section with
central strand of collapsed cells, cortical
cells large, thin-walled in 2-3 rows. Leaves
bract-like below, erect to spreading above;
ovate to ovate-lanceolate, acuminate, 0,9-1 ,8
mm long; margins plane, entire. Costa
excurrent as hair-point; in section round,
cells undifferentiated, substereids to stereids.
Laminal cells short-rhombic to rhomboidal;
marginal cells shorter or rectangular to
quadrate; basal cells short-rectangular to
quadrate.
Perichaetia terminal. Leaves erect, ovate-
lanceolate, acuminate, 1-2 mm long; margins
entire, plane to recurved. Costa excurrent as
hair-point; in section round, cells undiffer-
entiated, incrassate or stereids. Laminal
cells rhombic to rhomboidal, 70-90 (100) pm
long; apical and marginal cells shorter; basal
cells rectangular to quadrate, frequently
hyaline. Spores 12-20 per capsule, rounded-
triangular to tetrahedral, 120-185 pm, smooth
or granulose, yellowish. Fig. 19: 1-9.
This plant has been collected in woodlands of
central, northern and eastern Transvaal, Zululand
and Natal and is also known from Zaire. Map 26.
Map 26. — • Archidium acanthophyllum
x Archidium julicaule
Vouchers: Magill 5004; Van Vuuren 1772;
Wager PRE-CHI 9.
Archidium acanthophyllum may be separated
from the very similar A. ohioense and A. microthecium
on the basis of the short-rhombic to rhomboidal
laminal cells, long-excurrent costa, ovate to ovate-
lanceolate leaves and short-acuminate apices. This
species is the most frequently collected species of
Archidium in Southern Africa.
76
Archidiaceae
Archidiaceae
77
5. Archidium subulatum C. Mull, in
Flora, Jena 71:7 (1888). Snider in J. Hattori
bot. Lab. 39: 144 (1975). Type: Cape, Cape
Town, Rehmann s.n., 1876 (S-PA, lecto.!),
vide Snider (1975).
Plants in dense tufts, yellow to yellow-
brown; terricolous. Stems erect or prostrate,
old stems stoloniferous, 4-15 mm long, simple
or branching by subperichaetial innovations,
central strand small, cells thin-walled, inner
cortical cells large, thin-walled, in 2 rows,
outer cortical cells small, in 1-2 rows. Leaves
erect-spreading; ovate-lanceolate to lanceo-
late, acuminate, 0,9-1, 3 mm long; margins
plane to recurved above, entire to serrulate
above. Costa percurrent to short-excurrent;
in section round, central cells substereids,
surface cells incrassate. Laminal cells laxly
rectangular to rhomboidal above; basal
cells rectangular to quadrate below, lax,
lower marginal cells short-rectangular to
quadrate.
Perichaetia terminal. Leaves erect; ob-
long-acuminate, 1,3-2, 4 mm long; base
ovate; margins recurved above, entire. Costa
percurrent to short-excurrent; in section
round, central cells incrassate or substereids,
surface cells incrassate. Laminal cells laxly
rectangular to rhomboidal, 80-110 gm long,
basal cells larger, laxly rectangular. Spores
8-60 per capsule, rounded to irregularly
angular or polyhedral, 150-170 pm, smooth
to granulose, yellow. Fig. 19: 10-16.
Endemic to Southern Africa, this species grows
on sandy soils in the fynbos biome of the south-
western Cape. Map 25.
Voucher: Bews 8421.
Archidium subulatum can be distinguished by its
laxly rectangular median leaf cells and the percurrent
costa.
6. Archidium muelleranum Snider in
Bryologist 78: 154 (1975), as A. muellerianum.
Type: Cape, Rehmann 388 (S-PA, holo.).
Plants loosely caespitose, yellow to
yellow-green ; terricolous. Stems 3-5 mm high,
branching by subperichaetial innovations; in
section central strand present, inner cortical
cells large, incrassate, in 1-3 rows, outer
cortical cells smaller, incrassate, in 1-3 rows.
Leaves bract-like below, larger above, erect-
spreading to spreading, concave; ovate-
lanceolate, acuminate, 0,9-1, 2 mm long;
margins plane to slightly recurved, entire.
Costa percurrent to short-excurrent; in
section semi-circular, ventral surface flat,
cells incrassate or stereids dorsally, ventral
cells larger. Laminal cells irregularly short-
rectangular to quadrate or trapezoidal ; basal
cells not differentiated or slightly larger.
Perichaetia terminal. Leaves erect to
erect-spreading, frequently secund, concave;
broadly ovate-lanceolate to oblong-ovate,
acuminate to subulate, 1, 4-2,0 mm long;
base broadly ovate to oblong or concave;
margins narrowly recurved above, entire.
Costa percurrent to slightly excurrent, in
section semi-circular, ventral surface flat,
dorsal cells incrassate or stereids, ventral
cells larger. Laminal cells irregularly rect-
angular to rhomboidal or trapezoidal, 40-80
pm long, larger below, basal cells irregularly
rectangular, thin-walled, hyaline. Spores 20-40
per capsule, irregularly rounded triangular
to tetrahedral, 149-185/tm, densely papillose,
yellow to yellow-brown. Fig. 19: 17-26.
This plant is endemic to the southwestern Cape
and grows on sandy soil. Map 23.
Vouchers: Magill 4101; Parker 24849 (BOL!);
Rehmann 427.
Archidium muelleranum can be distinguished by
its densely papillose spores and strongly recurved
upper margins of the perichaetial leaves.
7. Archidium rehmannii Mitt, in J. Linn.
Soc., Bot. 22: 300 (1885); Snider in J. Hattori
bot. Lab. 39: 148 (1975). Type: Cape, Cape
Town, Rehmann s.n. (NY, holo.).
Fig. 19. — Archidium acanthophyllum (1-9): 1. habit, x 1 ; 2. habit, x 10; 3. stem in cross section, X 170; 4.
leaves, x40; 5. leaf apex, x 170; 6. perichaetial leaf, x40; 7. perichaetial leaf in cross section, x 170; 8. peri-
chaetial leaf cells, x435; 9. spore, x 170. A. subulatum (10—1 6) : 10. habit, x 1 ; 11. habit, x 10; 12-13. leaves,
x40; 14. perichaetial leaves, x40; 15. perichaetial leaf cells (left side), xl70; 16. cells in upper perichaetial
leaf, xl70. A. muelleranum (17-26): 17. habit, xl; 18. habit, xlO; 19. stem in cross section, xl70; 20-21.
leaves, x40; 22. perichaetial leaf, x40; 23. perichaetial leaf in cross section, x 170; 24. perichaetial leaf cells,
X435; 25. cells in upper perichaetial leaf, x 170; 26. spore, x 170. A. rehmannii (27-33): 27. habit, x 1 ; 28. habit,
x 10; 29. leaves, x40; 30. perichaetial leaf, x40; 31. perichaetial leaf in cross section, x435; 32. cells in upper
perichaetial leaf (right side), Xl70; 33. spore, Xl70. (1-9, Van Vuuren 1735; 10-16, Bews 8421; 17-26,
Rehmann 429 d; 27-33, Rehmann 427).
78
Archidiaceae
Plants caespitose, yellow-green; terri-
colous. Stems 2-10 mm high, branching by
subperichaetial innovations; in section with
central strand of collapsed cells, inner corti-
cal cells large, in 2 rows, outer cortical cells
smaller, in 1-2 rows. Leaves bract-like below,
larger above, erect to spreading; ovate-
lanceolate to lanceolate, acuminate, 0,9-1, 2
mm long, ventral surface flat to channelled;
base often clasping; margins plane to recurved
above, entire; lamina bistratose juxtacostally.
Costa percurrent; in section crescent-shaped,
ventral surface flat, cells undifferentiated,
incrassate. Lamina! cells irregular, trapezoi-
dal, short-rectangular, quadrate and short-
rhomboidal; basal cells irregularly short-
rectangular to quadrate.
Perichaetia terminal. Leaves erect, loosely
secund, concave, narrowly acuminate to
subulate above an ovate to oblong base, 1 ,2-
1,8 mm long; margins plane to recurved
above, entire to crenulate; lamina bistratose
juxtacostally. Costa percurrent to excurrent;
Map 27. — • Archidium rehmannii
X Archidium andersonianum
in section crescent-shaped, ventral surface
flat, cells undifferentiated, incrassate. Lamina!
cells irregular, rectangular to trapezoidal or
rhomboidal, 30-60 pm long, larger below;
basal cells irregularly rectangular to rhom-
boidal, thin-walled, hyaline. Spores 12-28 per
capsule, 132-167 pm, smooth to granulose,
yellow. Fig. 19: 27-33.
This species is endemic to the fynbos biome of
the southwestern Cape. Map 27.
Voucher: Rehmann 429.
The perichaetial leaves of A. rehmannii are
narrowly acuminate to long-subulate from an ovate
to oblong or occasionally elliptic base, the leaf
margins are plane or rarely recurved and the leaves
appear loosely secund. These characters define the
species.
8. Archidium capense Hornsch. in Lin-
naea 15: 135 (1841); Sim, Bryo. S. Afr. 141
(1926); Snider in Bryologist 78: 148 (1975).
Type: Cape, Ecklon, 1827 (H, lecto.), vide
Snider (1975).
Archidium campylopodium C. Mull, in Hedwigia
38: 52 (1899). Type: Cape Town, Rehmann s.n.,
1875 (S-PA, lecto.).
Archidium compactum fo. tenerior C. Mull, in
Hedwigia 38: 52 (1899), nom. nud. Type: Orange
Free State, Bloemfontein, Rehmann s.n.
Plants loosely caespitose, yellow to
yellow-green; terricolous. Stems 1-4 mm high,
simple or branching by subperichaetial
innovations; in section round, central strand
of collapsed cells, cortical cells large, incras-
sate, outer 1-2 rows smaller. Leaves bract-
like below, larger above, erect; ovate to
ovate-lanceolate, 0, 6-1,0 mm long; apex
acute or apiculate; margins plane, entire to
crenulate. Costa percurrent to apiculate; in
section semi-circular or crescent-shaped,
ventral surface flat, cells undifferentiated,
substereids or stereids. Laminal cells irregular,
short-rectangular to quadrate or trapezoidal ;
basal cells not differentiated.
Fig. 20. — Archidium capense (1-10): 1. habit, xl; 2. habit, xlO; 3. leaf, x40; 4. leaf apex (left side),
xl70; 5-6. innovation leaves, x40; 7. perichaetial leaves, x40; 8. perichaetial leaf cells (left side), xl70; 9.
cells in upper perichaetial leaf, xl70; 10. spore, xl70. A. julicaule (11-17): 11. habit, xl; 12. habit, xlO;
13. leaves, x40; 14. innovation leaf, X40; 15. perichaetial leaves, x40; 16. perichaetial leaf cells, x435; 17.
cells in upper perichaetial leaf (right side), x 170. A. amplexicaule (18-25): 18. habit, xl; 19. habit, x 10; 20.
leaves, x 30; 21. perichaetial leaves, x 30; 22. perichaetial leaf in cross section, x435; 23. perichaetial leaf cells,
X 170; 24. perichaetial leaf apex (right side), x 170; 25. spore, X 170. A. andersonianum (26—35) : 26. habit, X 1 ;
27. habit, x 10; 28. innovation leaf, x30; 29. leaves, x30; 30. perichaetial leaves, X30; 31. perichaetial leaf in
cross section, x 435; 32. basal cells of perichaetial leaf, x 170; 33. perichaetial leaf cells, x435; 34. perichaetial
leaf apex, x 170; 35. spore, xl70. (1-10, Magill 6326; 11-17, Rehmann 426; 18-25, Lorentz 1877; 26-35,
Almborn 5348).
Archidiaceae
79
80
Archidiaceae
Perichaetia terminal. Leaves erect to
erect-spreading; ovate to oblong-ovate or
elliptic, acuminate to subulate, 0,8-1, 7 mm
long; margins plane to weakly recurved
above, irregularly crenulate, frequently hya-
line above. Costa excurrent as hair-point; in
section semi-circular or crescent-shaped,
ventral surface flat, cells undifferentiated,
substereids to stereids. Laminal cells irregu-
lar, rhomboidal to fusiform and trapezoidal,
sometimes quadrate, 20-45 pm long, incras-
sate; basal cells irregularly rectangular, thin-
walled, hyaline in marginal region. Spores
8-40 per capsule, irregularly rounded to
polyhedral, 125-176 //m, smooth, pale yellow
to yellow-orange. Fig. 20: 1-10.
This plant has been collected in the southwestern
Cape, Sani Pass area of Lesotho and at Bloemfontein
in the Orange Free State. Map 28.
Map 28. — • Archidium capense
x Archidium amplexicaule
Vouchers: Magill 4384, 4410, 6326.
The perichaetial leaves with plane, irregularly
crenulate margins and costa excurrent as hair-point
help to define this species. The leaves are frequently
fragile and hyaline above.
9. Archidium julicaule C. Mull, in Hed-
wigia 38: 52 (1899); Snider in J. Hattori bot.
Lab. 39: 153 (1975). Type: Cape, Cape Town,
Rehmann 426 (S-PA, lecto.; PRE!), vide
Snider (1975).
Plants loosely caespitose, yellow to
yellow-green ; terricolous. Stems 2-7 mm high,
branching by subperichaetial innovations; in
section central strand large, cells small,
thin-walled, inner cortical cells in 2 rows,
incrassate, outer cortical cells smaller, in
1-2 rows, incrassate. Leaves bract-like below,
larger above, appressed; oblong-ovate, ob-
long or elliptical, (0,2-) 0,4 (-0,7) mm
long; apex acute or mucronate; margins
plane to slightly recurved below, irregularly
denticulate, teeth single, upper lamina bistra-
tose juxtacostally. Costa mucronate to short-
excurrent; in section crescent-shaped, ventral
surface flat, cells undifferentiated, incrassate
or stereids. Laminal cells irregularly short-
rectangular, rhombic, and short-rhomboidal,
incrassate; basal cells rectangular, thin-
walled to incrassate.
Perichaetia terminal. Leaves erect, con-
cave ; rounded-ovate or oblong-ovate, acumi-
nate, 1 ,0-1,4 mm long; margins plane,
entire or crenulate to irregularly denticulate.
Costa percurrent to short-excurrent; in
section semi-circular or crescent-shaped,
ventral surface flat, cells undifferentiated,
incrassate or stereids. Laminal cells irregularly
short-rectangular to quadrate and rhombic
or rhomboidal, 15-27 pm long, thin-walled
to incrassate; basal cells rectangular to
rhomboidal, thinner-walled, basal margins
hyaline. Spores 16-36 per capsule, angular to
polyhedral, 140-220 pm, smooth, yellowish.
Fig. 20; 11-17.
This species is represented by a few specimens
collected on sandy soil in the fynbos biome of the
Cape Peninsula. The Rehmann collection 429b is
apparently mixed since duplicates at several institu-
tions have been given various names. Map 26.
Voucher: Type only.
The appressed leaves with irregularly denticulate
margins and acute apices will help to identify this
species.
10. Archidium amplexicaule C. Mull, in
Linnaea 43: 346 (1881); Snider in J. Hattori
bot. Lab. 39: 151 (1975). Type: Concepcion
del Uruguay, Lorentz 1877 (S-PA, lecto.!),
vide Snider (1975).
Archidium chrysosporum Schimp. ex Jaeg. in Verh.
St Gall, naturw. Ges. 1868-69: 67 (1869), nom.
nud. Type: Cape, Saldanha Bay, sin. coll. (S-PA).
Plants gregarious or in tufts, yellowish;
terricolous. Stem erect, 3-10 mm high,
branching by subperichaetial, julaceous inno-
vations; in section central strand small,
cortical cells undifferentiated, in 3 rows.
Leaves bract-like, distant below, crowded
above, appressed to erect-spreading; ovate
Archidiaceae
81
to ovate-lanceolate, acuminate, 0,8-1, 2 mm
long; apex occasionally apiculate; margins
plane or narrowly recurved, entire. Costa
weak, percurrent. Laminal cells irregular,
quadrate to rectangular or rhomboidal;
basal cells rectangular.
Perichaetia terminal. Leaves erect; ovate
to oblong-acuminate, 1,0-1, 8 mm long,
margins plane to narrowly recurved above,
entire to slightly crenulate above. Costa weak,
percurrent to short-excurrent, in section semi-
circular, cells undifferentiated. Laminal cells
irregular, rectangular and rhombic to rhom-
boidal, 23-36 pm long; basal cells irregularly
rectangular, thin-walled, hyaline. Spores 8-36
per capsule, irregularly rounded to tetra-
hedral, 130-210 pm, smooth, yellowish. Fig.
20: 18-25.
This species occurs in South America and in the
Flora area it is restricted to the southwestern Cape.
Map 28.
Voucher: Type only.
Archidium amplexicaule is closely related to A.
andersonianum; but is separated by its julaceous
innovations, smaller ovate to oblong-ovate peri-
chaetial leaves and percurrent costae.
1 1 . Archidium andersonianum Snider in
Bryologist 78: 158 (1975). Type: Cape,
Stellenbosch Flats, Almborn 5348 (S-PA,
holo. !).
Plants loosely caespitose, yellow-green;
terricolous. Stem erect, 2-6 mm high, branch-
ing by subperichaetial innovations, in section
inner cortical cells in 1-2 rows. Leaves bract-
like below, larger above, erect to spreading,
concave; ovate to oblong, acuminate, 1,1-
1,3 mm long; margins plane to occasionally
narrowly recurved above, entire to crenulate.
Costa strong-excurrent as short hair-point;
in section semi-circular, ventral surface
flat, cells incrassate or stereids. Laminal
cells irregular, fusiform to rhomboidal; basal
cells rectangular.
Perichaetia terminal. Leaves erect, con-
cave; broadly oblong-ovate, acuminate, 1,3-
1,8 mm long; margins plane to narrowly
recurved, entire to infrequently crenulate.
Costa apiculate to excurrent as hair-point; in
section semi-circular to crescent-shaped,
ventral surface flat, ventral cells incrassate,
dorsal cells smaller, incrassate or stereids.
Laminal cells irregular, rhomboidal to fusi-
form, 50-75 pm long; rectangular at margin;
basal cells irregularly rectangular to rhom-
boidal, hyaline in marginal region. Spores
12-52 per capsule, irregularly rounded-
tetrahedral to polyhedral, 150-200 pm,
smooth, yellow. Fig. 20: 26-35.
Endemic to the southwestern Cape, this species
is represented by a single specimen, collected on
sandy soil on the Cape Flats. Map 27.
Voucher: Type only.
Although similar in habit to A. capense and A.
muelleranum, this species differs in the deltoid inno-
vation leaves with costa ending in a strong hair-point
and the larger, broadly oblong-ovate perichaetial
leaves with narrower costae.
Insufficiently Known Species
Archidium laterale Bruch ex Krauss in Flora,
Jena 29: 132 (1846). Type: Natal, Umslutie River,
Krauss s.n. Sim (1926) placed this species in the
synonomy of the Southern African species presently
treated as A. rehmannii Mitt., although he did not see
the type. Snider (1975), in his revision of Archidium,
was unable to locate the type of A. laterale and it has
not been found during this study.
83
DITRICHACEAE
Plants minute to medium-sized, in dense tufts or gregarious; terrestrial. Stems simple or
branched, central strand present or absent. Leaves ovate, oval or oblong, acute to subulate.
Costa strong, narrow or broad; in section round or flattened, generally with stereid bands.
Laminal cells generally rectangular, smooth or occasionally papillose above; basal cells
generally larger, rectangular; alar cells not distinct.
Seta straight or cygneous ; capsule cleistocarpic or stegocarpic, long-exserted or immersed ;
peristome single or absent, teeth 16, fragile, cleft or irregularly perforated, papillose or striate,
reddish; operculum rostrate; calyptra cucullate or mitriform.
The family comprises 19 genera of which 7 are known from Southern Africa.
1 Leaves distichous or tristichous:
2 Leaves distichous, flexuose 6. Distichium
2 Leaves tristichous, rigidly appressed 7. Tristichium
1 Leaves not in distinct rows:
3 Plants small to minute, gregarious or in small groups; capsule immersed or exserted
laterally on arcuate seta; peristome absent:
4 Plants small; leaves appressed to erect-spreading; capsule immersed; seta straight
1. Pleuridium
4 Plants minute; leaves secund; capsule exserted laterally by arcuate seta:
5 Leaf cells smooth; capsule stegocarpic; calyptra mitriform 2. Eccremidium
5 Upper leaf cells papillose; capsule cleistocarpic; calyptra cucullate 1. Pleuridium
3 Plants small to medium, in dense tufts or gregarious; capsule long-exserted; peristome
present :
6 Plants glaucous; branched above; gregarious in caves and deep crevices. ...5. Saelauia
6 Plants dark green to yellowish green, infrequently branched, forming dense tufts or
patches in open sites:
7 Leaves short, apex acute; leaf cells quadrate; capsule curved, plicate when dry
4. Ceratodon
7 Leaves long, apex acuminate to subulate; leaf cells elliptical to rectangular;
capsule straight, smooth when dry 3. Ditrichum
1. PLEURIDIUM
Pleuridium Rabenh., Deutschl. Kryptogamenfl. 2: 79 (1848), nom. cons.; Broth, in Natiirl.
PflFam. 10: 157 (1924); Sim, Bryo. S. Afr. 142 (1926). Lectotype species: P. subulatum (Hedw.)
Rabenh., vide Snider & Margadant in Taxon 22: 691 (1973).
Plants minute to small, gregarious, yellowish green; terricolous. Stems erect, 1-3 mm tall,
to 10 mm in sterile specimens. Leaves erect or patent, small, oval to ovate, acute to long-
acuminate above. Costa percurrent. Laminal cells quadrate to rectangular, smooth or papil-
lose.
Autoicous. Perichaetia terminal, leaves larger. Capsule cleistocarpic, sessile or laterally
exserted, urn round to elliptical, exothecial cells distinct; spores papillose or spinose.
Although the 34 species of Pleuridium are generally restricted in distribution, the genus is known from all
continents except Antarctica. The four Southern African species are endemic and have very restricted ranges.
84
Ditrichaceae
1 Plants elongated, 5-10 mm tall; stems in section fluted, without central strand; leaf cells frequently prorate
(obvious only in cross section) 3. P. ecklonii
1 Plants small to minute, to 3 mm tall; stems in section entire, central strand present; leaf cells smooth or
papillose:
2 Plants minute; upper leaf cells with several low, blunt papillae over lumen 4. P. papillosum
2 Plants small; upper leaf cells smooth:
3 Leaves patent, apices acuminate from near stem base; capsule globose 1. P. pappeanum
3 Leaves appressed, apices shorter below, gradually longer distally, capsule oval to elliptical. .2. P. nervosum
1. Pleuridium pappeanum (C. Mull.)
Jaeg. in Verh. St Gall, naturw. Ges. 1871-72;
373 (1872); Broth, in Natiirl. PflFam. 10:
157 (1924); Sim, Bryo. S. Afr. 143 (1926).
Syntypes: Cape, Swellendam, Pappe s.n. ;
Swellendam, Ecklon s.n. (BM!).
Astomum pappeanum C. Miill., Syn. Muse. 1: 15
(1848).
Astomum breutelianum Hampe ex C. Miill. in
Bot. Ztg 17: 197 (1859). Pleuridium breutelianum
(Hampe) Jaeg. in Verh. St Gall, naturw. Ges. 1871-72:
374 (1872); Broth, in Natiirl. PflFam. 10: 157
(1924). Type: Cape, Gnadenthal, Breutel s.n. (BM!).
Bruchia rehmannii C. Miill. in Flora, Jena 71: 10
(1 888). Sporledera rehmannii (C. Miill.) Kindb.,
Enum. Bryin. Exot. 95 (1889); Broth, in Natiirl.
PflFam. 10: 158 (1924). Type: Cape, Rondebosch,
Rehmann s.n., Aug. 1875 (NH!).
Plants small, gregarious, light to yellow-
ish green; terricolous. Stems erect, to 3 mm
tall, simple or branched below; in section
with small central strand, cortical cells
large, thin-walled, outer 1-2 rows slightly
smaller, reddish brown. Leaves patent, lower
leaves oval, 0,6-0, 8 mm long; upper leaves
oval to oblong, abruptly subulate, 1-2 mm
long; apex acute; margins erect, entire.
Costa percurrent; in proximal section guide
cells exposed ventrally, dorsal cells in 2-3
rows, incrassate; in distal section guide cells
2-4, thickened, ventral cells in 2 rows, sub-
stereids, dorsal substereid band 2-3 cells
thick, dorsal surface cells slightly larger.
Laminal cells of lower leaves short-rectangu-
lar; in upper leaves linear above shoulders;
basal cells long-rectangular.
Autoicous. Perigonium bud-like, axillary.
Perichaetial leaves oval, gradually long-
acuminate, 3, 0-3, 2 mm long; margins
entire; costa percurrent; laminal cells linear
above, oblong-hexagonal at shoulders; basal
cells rectangular. Seta short, 0, 1-0,2 mm
long; capsule emergent, urn globose, 0,6-0, 8
mm long, orange, very short-rostrate, exothe-
cial cells quadrate to rectangular; calyptra
cucullate; spores round, 17-20 pm, sparsely
papillose. Fig. 21 : 16-26.
Map 29. — • Pleuridium pappeanum
x Pleuridium papillosum
Fig. 21. — Pleuridium nervosum (1-15): 1. habit, x 1 ; 2. habit, x 10; 3-5. leaves, x40; 6. leaf in proximal
cross section, x 320; 7. leaf in distal cross section, x 320; 8. cells at basal margin, x 320; 9. cells at upper margin,
X320; 10. cells at upper margin of lower leaf, x320; 11. laminal cells of perichaetial leaf, x320; 12. peri-
chaetial leaf, x20; 13. calyptra, x50; 14. exothecial cells at capsule apex and spores, x 50; 15. spore, x240.
P. pappeanum (16-26): 16. habit, X 1 ; 17. habit, x 10; 18. stem in cross section, x 150; 19. lower stem leaf, x40;
20. upper stem leaf, x40; 21. basal leaf cells, x320; 22. laminal cells, x320; 23. marginal cells at base of
subula, x320; 24. cells in leaf subula, x320; 25. perigonium, x50; 26. exothecial cells at capsule apex, x50.
P. ecklonii (27-34): 27. habit, x 1 ; 28. habit, x2; 29. stem in cross section, X 150; 30. leaves, x55; 31. leaf in
cross section, x425; 32. basal leaf cells, x320; 33. leaf cells at base of subula, x320; 34. leaf apex, x435.
(1 2, Scheipe 7785; 3-15, Mag ill & Schelpe 3987; 16-26, Magill & Schelpe 3920; 27-30, Sim 9259; 31-34, Wager
20).
Ditrichaceae
85
86
Ditrichaceae
Endemic to Southern Africa, P. pappeanum is
infrequently found on thin sandy soils under brush
in the western Cape. Recent collections have been
made in the arid shrublands of Namaqualand, far
north of the original collection sites near Swellen-
dam. Map 29.
Vouchers: Magill & Schelpe 3820, 3859.
The abrupt change from acute to long-acuminate
apices of the patent leaves on the lower stem and the
globose capsule will define this species.
2. Pleuridium nervosum {Hook.) Mitt, in
Hooker, J. Bot. Kew Gdn Misc. 8: 257
(1856); Broth, in Naturl. PflFam. 10: 157
(1924); Sim, Bryo. S. Afr. 15: 142 (1926).
Type: Cape, Menzies s.n., 1791 (BM, holo.!).
Phascum nervosum Hook., Musci Exot. 2: 105
(1819). Astomum nervosum (Hook.) C. Miill. in Bot.
Ztg 5: 98 (1847).
Plants small, in scattered groups, yellow-
ish green ; terricolous. Stems erect, julaceous,
to 3 mm tall, simple; in section with small
central strand, cortical cells incrassate,
margin lobed. Leaves imbricate, lower leaves
ovate to oval, 0,4-0, 8 mm long, upper
leaves oblong, to 1,2 mm long; apex obtuse
to acute in lower leaves, gradually longer
pointed to acuminate in upper leaves;
margins irregularly serrate by projecting
upper and/or lower cell ends. Costa per-
current in lower leaves, filling acumen in
upper leaves; in section guide cells 5-8, large,
ventrally exposed, dorsal stereid band small,
dorsal surface cells large, strongly incrassate.
Upper laminal cells of lower leaves sub-
quadrate to short-rectangular becoming
rhombic to rhomboidal in subperichaetial
leaves; basal cells quadrate to rectangular in
all leaves.
Autoicous. Perichaetial leaves broadly
oval, abruptly short-acuminate, 2,2-2, 5 mm
long; margins serrulate at shoulders; costa
percurrent; laminal cells rhombic to rhom-
boidal; marginal cells generally narrower,
basal cells rectangular. Seta short, to 0,4
mm long; capsule immersed, urn oval to
elliptical, 0,8 mm long, blackish red to
yellowish red, obtuse to very shortly beaked,
exothecial cells longitudinally short-rectangu-
lar; calyptra cucullate; spores round to
triangular, 20-25 pm, papillose. Fig. 21:
1-15.
Known from Australia, New Zealand and
Southern Africa, P. nervosum inhabits dry, open
soils or shaded areas under brush, in the western,
central and eastern Cape and Natal. In the western
Cape, plants are frequently collected in association
with P. pappeanum. Map 30.
Vouchers: Giffen PRE-CH4388; Magill &
Schelpe 3987; Smook 1120.
This species is separated from the other species
of Pleuridium by its julaceous stems, leaf apices
gradually changing up the stem from acute to acumi-
nate and the oval to short-oblong capsule. The spores
are consistently larger and less highly ornamented
than those of P. pappeanum.
3. Pleuridium ecklonii ( C . Miill.) Snider
in J. Hattori bot. Lab. 39: 155 (1975).
Type: Cape, Cape Town, Naumann, 1874
(BM, lecto.), vide Snider (1975).
Archidium ecklonii Hampe ex C. Miill. in Hedwigia
38: 53 (1899); Broth, in Naturl. PflFam. 1 : 289 (1909);
Sim, Bryo. S. Afr. 141 (1926).
Plants slender, forming intertwining
tufts, green; terricolous. Stems erect, 5-10
mm high, simple, with leaves distant; in
section without central strand, inner cortex
of large, thin-walled cells, outer cortical cells
in 2-3 rows, small, incrassate, reddish, margin
fluted. Leaves patent, oval-acute to ovate-
acuminate, 0, 5-1,0 mm long; margins
entire, recurved below. Costa percurrent,
reflexed at tip of oval-acute leaves ; in section
guide cells 4, large, ventral cells incrassate,
dorsal substereid band in single layer, dorsal
surface cells incrassate. Laminal cells quad-
rate to rectangular, frequently weakly
prorate ; basal cells larger, rectangular.
Sporophytes have not been found;
Snider (1975) describes the perichaetial leaves
as abruptly linear-subulate from a short-oval,
sheathing base, one or two rows of marginal
cells of the leaf base are narrower and more
or less differentiated from marginal cells,
the basal cells are faintly yellow-orange, and
the lowermost row of basal cells is slightly
inflated. The costa has guide cells and stereid
cells in transection. Fig. 21 : 27-34.
Fig. 22. — Pleuridium papillosum (1-12): 1. habit, xl; 2. habit, xlO; 3. habit, x25; 4-6. leaves, x40;
7. leaf in cross section, x 435; 8. basal leaf cells, x 435; 9. leaf apex, x 435; 10. perichaetial leaf, x 40; 11. capsule
X 40; 12. spore, x 380. Eccremidium exiguum (13-26): 13. habit, x 1 ; 14. habit, x 10; 15. stem in cross section,
x 240; 16-18. leaves, x 40; 19. leaf in cross section, x 435; 20. basal leaf cells, x 170; 21. cells at base of leaf
subula, x 170; 22. leaf apex, x 170; 23. perichaetial leaf, x40; 24. sporophyte and perichaetial leaves, x40;
25. calyptra, X 60; 26. capsule, x 60. (1-12, Schelpe 7624; 13-26, Stone s.n.).
,o-Pj2lo
Ditrichaceae
87
88
Ditrichaceae
This species has been occasionally collected in
and around Cape Town since its original discovery by
Breutel in 1858 and Naumann in 1874. A specimen
collected by Wager in 1911 from Van Reenen, north-
western Natal, has been verified, but it presents an
unusual disjunction and may be in error. Map 30.
Vouchers: Pillans 4073; Rehmann 425; Sim
9259; Wager 85.
Map 30. — • Pleuridium nervosum
x Pleuridium ecklonii
The plants are quite distinct from other South
African species. The elongated, fluted stems and small
distant leaves easily divide the sterile specimens from
P. pappeanum and P. nervosum. The change in leaf
apices up the stem from acute to acuminate and back
again, indicates several seasons’ growth. Add to this
stem anatomy and absence of sporophytic characters
and it becomes clear that the present generic place-
ment is doubtful. Until fertile material can be found,
however, nothing can be done to solve the problem.
4. Pleuridium papillosum Magill, sp. nov.,
P. arnoldii ( R . Br. ter.) Par. foliis falcato-
secundis setaque arcuata simile, sed cellulis
papillosis subulae et sporis majoribus differt.
Type; Cape, 11 km S of Clanwilliam on
road to Algeria Forest Station, on clay bank.
Schelpe 7624 (BOL, holo.; CINN; MO;
PRE).
Plants minute, gregarious, yellowish
green; terricolous. Stems very short, to 0,5
mm long, simple; in section round, central
strand present, cortical cells large, in 3 rows,
outer row reddish. Leaves spreading; lower
leaves small, ovate-acute, 0, 5-1,0 mm;
upper and subperichaetial leaves oblong-
acuminate, 1,0-1, 8 mm long; apex reflected
above clasping base; margins plane, crenu-
late above shoulders. Costa percurrent; in
proximal section flattened, consisting of 5-6
substereid cells in 2-3 rows; in distal section
rounded, of 8-10 substereid cells in 3 rows.
Laminal cells rectangular in subula, papillose ;
elongate-hexagonal to rectangular at should-
ers, increasing in size toward leaf base; in
section gradually decreasing in size toward
margin.
Perigonia not seen. Perichaetia terminal ;
leaves falcate-secund, linear-subulate, con-
volute, 2, 2-3,0 mm long. Seta 0,5-0, 6 mm
long, yellow, arcuate in opposite direction to
perichaetial leaves; capsule cleistocarpic,
laterally exserted; urn ovoid, short-rostrate,
0,8 mm long, reddish yellow; exothecial
cells quadrate proximally and distally, median
cells rectangular; stomata present around
capsule base; calyptra cucullate, small,
covering upper capsule only, long-beaked,
0,5 mm long; spores round, 35-38 pm,
spinose. Fig. 22: 1-12.
Endemic to South Africa. The only collection,
so far known, comes from a clay road cutting south
of Clanwilliam along the dirt road to Algeria Forest
Station. Map 29.
Voucher: Type only.
The species is identified by its narrow leaves
with clasping base and reflexed, papillose subula, and
its laterally exserted capsule and large, sp inose spores.
2. ECCREMIDIUM
Eccremidium Hook. & Wils. in Hooker’s, Lond. J. Bot. 5: 450(1846); Broth, in Natiirl. PflFam.
10; 159 (1924); Scott & Stone, Moss. S. Aust. 125 (1976). Type species: Not designated.
Plants small to minute, gregarious; terricolous in open areas. Stems erect; central strand
present. Leaves secund, oval-subulate; costa filling subula. Laminal cells oblong, smooth.
Monoicous. Seta short, curved; capsule operculate, laterally exserted, gymnostomous;
calyptra mitriform.
Eccremidium has 6 species, 5 endemic to Australia, the sixth, E. exiguum , is found in Australia and South
Africa.
Ditrichaceae
89
Eccremidium exiguum (Hook, f & Wils.)
Wils. ex Salm. in Rev. Bryol. 27: 85 (1900);
Broth, in Natiirl. PflFam. 10: 159 (1924);
Scott & Stone, Moss. S. Aust. 125 (1976).
Type: Australia, Swan River, Drummond
s.n. (BM).
Phascum exiguum Hook. f. & Wils. in Hooker’s
Icon. PI. 8:737 B (1848).
Plants very small to minute, gregarious,
light green; terricolous. Stems erect, 0,5 mm
tall; in section round, central strand present,
cortical cells very thin-walled, in 3-4 rows,
outer row weakly thickened, reddish. Leaves
secund; oval, long-subulate, 1,3-1, 7 mm
long; margins plane, entire. Costa filling
subula; in section ventral cells thin-walled,
in single layer, dorsal cells in 2 layers, cells
smaller, incrassate. Laminal cells rectangular,
thin-walled, cells of subula linear, incrassate,
smooth.
Paroicous. Perichaetia terminal, leaves
undifferentiated, 2,2 mm long. Seta arcuate,
0,8 mm long; capsule laterally exserted,
pendent, ovoid, 0,8 mm long (immature),
exothecial cells rounded-quadrate, incras-
sate, transversely elongate at mouth; peri-
stome absent; operculum short-rostrate;
calyptra mitriform; spores not seen. Fig. 22:
13-26.
Map 31. — • Ceratodon purpureus
x Eccremidium exiguum
The species, previously known only from Austra-
lia, was recently collected in the northern Drakens-
berg, at Royal Natal National Park. Map 31.
Voucher: Stone s.n., 23 Mar. 1972 (BOL!;
MELU).
The small plants with secund leaves and pendent,
operculate capsule are very distinctive. The plants
could perhaps be confused with Pleuridium papillo-
sum, but there the upper leaf cells are papillose and
the capsules cleistocarpic.
3. DITRICHUM
Ditrichum Hampe'm Flora, Jena 50: 181 (1867), nom. cons.; Broth, in Natiirl. PflFam. 10: 161
(1924); Sim, Bryo. S. Afr. 145 (1926). Type species: D. homomallum (Hedw.) Hampe.
Plants small to large, in tufts, yellow-green to olive-green; terricolous. Stems erect,
infrequently branched, with central strand. Leaves flexuose, narrow, acuminate to setaceous.
Costa subpercurrent to excurrent, ventral stereid band present or absent. Laminal cells rec-
tangular to quadrate, incrassate.
Autoicous. Perigonia gemmate, axillary; perichaetia terminal. Seta straight, long;
capsule erect, elliptical or cylindrical, generally symmetrical; peristome teeth 16, lanceolate,
divided to base or irregularly cleft or perforated, papillose; operculum rostrate; calyptra
cucullate.
Ditrichum is a widespread genus with 87 species. There appears to be no concentration of species, each
major continental area, except Antarctica, has 12-15 species. Sterile plants may be confused with Dicranella
or other genera of the Dicranaceae, however the costal anatomy should identify Ditrichum. The cylindrical,
non-furrowed capsule and papillose peristome will place fertile plants. Within Ditrichaceae, the long leaves,
placed evenly around the stem, and rectangular cells of the long leaf subula, will define the genus.
Four species are presently recognized in Southern Africa. Exclusion of D. amoenum (Thwait. & Mitt.) Par.
from the Flora and the placement of D. hymenodontium Dix. and D. spirale Dix. into synonomy were necessary
because of the variability expressed by the gametophytes and the fragile nature of the peristome teeth.
90
Ditrichaceae
1 Cells of subula short-rectangular to elliptical:
2 Leaves 5, 0-6, 5 mm long; subula strongly twisted when dry 2. D. punctulatum
2 Leaves 2, 5-4, 5 mm long; subula not twisted when dry 1. D. strictum
1 Cells of subula rectangular to linear:
3 Seta short, 5-7 mm; capsule short-cylindrical, 1,0-1, 5 mm long; leaves 1,0-2, 5 mm long, patent
3. D. brachypodum
3 Seta long, 10-20 mm; capsule cylindrical, 2, 5-3, 5 mm long; leaves 2, 0-3, 5 mm long, flexuose
4. D. difficile
1 . Ditrichum strictum {Hook. f. & Wils.)
Hampe in Flora, Jena 50; 182 (1867); Broth,
in Natiirl. PflFam. 10: 162 (1924); Sim, Bryo.
S. Afr. 147 (1926). Syntypes: Lord Auck-
land’s group, ? Hooker, Campbell’s Island,
Menzies (BM !).
Lophiodon strict us Hook. f. & Wils. in Hooker,
Lond. J. Bot. 3: 544(1844).
Plants small to medium, in dense tufts,
green to yellow-green above, brownish below;
terricolous. Stems 20-30 mm high, infre-
quently branched, radiculose below; in
section round, central strand small, cortical
cells in 3-4 rows, outer row smaller, incras-
sate. Leaves erect to patent; subulate above
oval base, 2, 5-4, 5 mm long; apex blunt,
toothed; margins erect, entire below, toothed
in upper subula; base weakly clasping,
gradually narrowing to subula. Costa broad,
filling subula; in proximal section sub-oval,
guide cells 6-10, ventral stereid band 2-3
cells thick, ventral surface cells larger, incras-
sate, dorsal stereid band 2-4 cells thick,
dorsal surface cells larger, incrassate or
substereids. Upper laminal cells incrassate,
elliptical to short-oblong in subula, rectangu-
lar at shoulders; basal cells long-rectangular,
strongly thickened; basal marginal cells
linear, thin-walled, hyaline.
Autoicous. Perichaetial leaves oblong-
subulate, 4 mm long. Seta erect, 5 mm long,
yellow; capsules oval to short-elliptical,
1, 5-2,0 mm long; peristome fragile, short,
Map 32. — • Ditrichum difficile
X Ditrichum strictum
teeth undivided, papillose; operculum ros-
trate, to 1 mm long ; calyptra and spores not
seen. Fig. 23: 21-27.
Although widespread on the subantarctic islands,
this species is rare in Southern Africa. In the Flora
area, D. strictum is known only from the Natal
Drakensberg, above 5 500 m. Map 32.
Vouchers: Sim 8538, 8685.
This species is identified, in Southern Africa,
by its short-elliptical upper leaf cells and straight,
erect subula. Ditrichum strictum is related to several
other species from New Zealand, Australia and the
subantarctic islands, e.g. D. punctulatum and D.
cylindricarpum (C. Mull.) F. Miill., but it lacks the
pronounced leaf shoulders or spirally twisted subula
exhibited by these species.
Fig. 23. — Ditrichum brachypodum (1-11): 1. habit, xl; 2. habit, x 10; 3. stem in cross section, x200; 4.
leaves, x40; 5. leaf in proximal cross section, x370; 6. leaf in distal cross section, x370; 7. basal leaf cells at
margin, x290; 8. upper laminal cells at margin, x290; 9. leaf apex, x290; 10. perigonium, x50; 11. part of
capsule mouth with peristome teeth and spores, x 167. D. punctulatum (12-20) : 12. habit, x 1 ; 13. part of stem,
x 10; 14. stem in cross section, x200; 15. leaf, x25; 16. leaf in mid-leaf cross section, x370; 17. leaf in distal
cross section, x370; 18. basal leaf cells at margin, x200; 19. upper laminal cells at margin, x200; 20. leaf
apices, x200. D. strictum (21-27): 21. habit, x 1; 22. part of stem, x 10; 23. leaves, x30; 24. leaf in cross
section, x 370; 25. basal leaf cells at margin, x 170; 26. laminal cells at shoulder, x 170; 27. leaf apex, x 170.
(1-3 & 10-11, Rehmann 86; 4-9, Cholnoky 520; 12-20, Henderson 232; 21-27, Sim 8538).
Ditrichaceae
91
92
Ditrichaceae
2. Ditrichum punctulatum Mitt, in Phil.
Trans. R. Soc. 168: 25 (1879); Broth, in
Natiirl. PflFam. 10: 162 (1924); Sainsb.,
N. Zeal. Mosses 76 (1955); Seppelt & Stone
in J. Bryol. 9: 321-325 (1977). Type: New
Zealand.
Ditrichum spirale Dix. in Trans. R. Soc. S. Afr. 8:
181 (1920); Sim, Bryo, S. Afr. 148 (1926). Type:
Cape, Gaika’s Kop, Tjumie, D., B. & M. Henderson
232 (BM, holo.!; PRE!).
Plants medium to large, loosely caespi-
tose, green to yellow-green; saxicolous.
Stems 30-70 mm high, infrequently branched;
in section central strand small, inner cortical
cells in 2-3 rows, large, weakly thickened,
outer cortical cells in 2 rows, smaller, strongly
incrassate. Leaves flexuose to falcate, secund
dry or wet; oval, abruptly or gradually
contracted to a long, spirally twisted subula,
4, 5-6, 5 mm long; apex acute, toothed;
margins erect to incurved, entire below,
dentate in upper subula. Costa broad, filling
subula; in proximal section flattened, guide
cells small, 10-12, dorsal and ventral stereid
bands 2-4 cells thick, surface cells not
differentiated or occasionally substereids,
laminal cells strongly thickened; in distal
section reniform, guide cells small, 4-6,
dorsal and ventral stereid bands 2-4 cells
thick, surface cells occasionally substereids.
Laminal cells variable in shape, elongate,
sinuolate, strongly incrassate, abruptly qua-
drate at shoulders; short, rounded-rectangu-
Map 33. — • Ditrichum brachypodum
X Ditrichum punctulatum
lar in lower subula; basal cells long-rectangu-
lar, thin-walled toward margin.
Sporophyte not known from Africa.
Fig. 23: 12-20.
Ditrichum punctulatum is known from Australia,
New Zealand and South Africa. In Southern Africa
the species has been collected in the Drakensberg of
Natal and on Gaika’s Kop in the eastern Cape. All
collections were made in subalpine grassland above
5 500 m. Map 33.
Vouchers: Esterhuysen 18620; Owen 10.
The very long, spirally twisted subula will help
to identify this species in Southern Africa. Variation
is seen in the abruptness with which the leaves con-
tract to the subula and in the extent of curvature of
the leaves. Ditrichum spirale is a smaller plant but
otherwise differs little from the typical form.
3. Ditrichum brachypodum (C. Mull.)
Broth, in Natiirl. PflFam. 1: 300 (1901);
Sim, Bryo. S. Afr. 145 (1926). Type: Orange
Free State, Kadziberg, Rehmann 86 (BOL!;
NH!;PRE!).
Leptotrichum brachypodum C. Mull, in Hedwigia
38: 89 (1899).
Leptotrichum brevifolium C. Miill. in Hedwigia
38: 88 (1899). Ditrichum brevifolium (C. Miill.)
Broth., horn, illeg., in Natiirl. PflFam. 1: 300 (1901),
non (Kindb.) Par. (1896). Type: Transvaal, near
Spitzkop, Wilms s.n., Apr. 1887 (G, holo.!).
Ditrichum hymenodontium Dix. in Trans. R. Soc.
S. Afr. 18: 249 (1930). Type: Transvaal, Benoni,
Wager 1005 (BM, holo.!).
Plants small, erect, yellowish green;
terricolous. Stems 2-5 mm high, simple; in
section round, central strand present, inner
cortical cells large, thin-walled, outer cortical
cells in 2 rows, stereids. Leaves patent,
lamina narrow, bistratose above; oval to
ovate, 1,0-2, 5 mm long; apex acuminate;
margin plane, entire to serrulate at apex.
Costa broad, occupying ^ to | of leaf width,
ending in acumen; in proximal section flat-
tened, guide cells 10, exposed ventrally,
large, incrassate, dorsal stereid band 1-2
cells thick, dorsal surface cells small, strongly
incrassate; in distal section dorsal and ventral
surface cells similar to laminal cells, incras-
sate, surrounding a single-layered median
stereid band. Laminal cells rectangular or
infrequently quadrate above, incrassate; basal
cells larger, narrow-rectangular.
Autoicous. Perichaetial leaves ovate-
acuminate, 2, 5-3, 5 mm long. Seta 5-7 mm
long, yellow, reddish with age; capsule erect,
Ditrichaceae
93
urn short-cylindrical, 1,0-1, 5 mm long;
peristome fragile, to 0,3 mm high, teeth
irregularly thickened, deeply cleft, frequently
with large perforations, papillose; operculum
rostrate, 0,4 mm long; calyptra cucullate,
2,5 mm long; spores round, 14-18 //m,
papillose. Fig. 23: 1-11.
Endemic to Southern Africa, D. brachypodum is
frequently collected in Natal and the northern, central
and eastern Transvaal. A few collections have also
been made in Swaziland, the Orange Free State, and
the eastern, southern and southwestern Cape.
Growing on soil, the species is frequently collected
along stream banks and in road cuttings. Map 33.
Vouchers: Crosby & Crosby 7495; Rodin 4532;
Sim 9751.
The overall size of D. brachypodum generally
separates it from the other larger species. The capsule
is short-cylindrical and the seta is less than 10 mm
long. The peristome is irregularly perforated, cleft
and thickened, and generally very fragile. The type
of D. hymenodontium consists of somewhat larger
plants with peristomes almost intact, but conforms
in all other characters to D. brachypodum. In Southern
Africa the distributions of D. difficile and D. brachy-
podum are sympatric and they are frequently collected
together in Natal. The former is more common in the
Cape, while the latter ranges primarily northward into
the Transvaal.
4. Ditrichum difficile (Dub.) Fleisch.,
Musci FI. Buitenzorg 1: 300 (1904); Scott &
Stone, Moss. S. Aust. 112 (1976). Type:
Java, Zollinger 411.
Trichostomum difficile Dub. in Moritzi, Syst.
Verz.Zoll. Pfl. 134(1846).
Ditrichum flexifolium Hampe in Flora, Jena 50: 182
(1867); Broth, in Natiirl. PflFam. 10: 162 (1924);
Sim, Bryo. S. Afr. 146 (1926). Type: Cape, Menzies
s.n., 1791 (BM, holo.!).
Plants small to medium, in tufts, yellow-
ish green; terricolous. Stems 7-12 mm high,
simple; in section round, central strand
present, inner cortical cells large, thin-walled,
outer cortical cells in 2-3 rows, substereids.
Leaves flexuose, oval to oblong, long-
subulate, 1 ,8-3,5 mm long; margins plane to
erect in subula, entire. Costa broad, filling
subula; in proximal section flattened, guide
cells 12-18, large, exposed ventrally or
frequently with 2-3 ventral surface cells
over central guide cells, dorsal stereid band
2-3 cells thick, dorsal surface cells not diffe-
rentiated; in distal section guide cells 18-20,
large, median, ventral stereid band 1-2 cells
thick, ventral surface cells substereids, dorsal
stereid band 2-4 cells thick, dorsal surface
cells substereids, dorsal surface irregular.
Laminal cells narrowly rectangular, incras-
sate; basal cells longer, oblong, incrassate.
Autoicous. Perichaetial leaves oblong,
abruptly setaceous, 5, 5-6, 5 mm long. Seta
10-20 mm long, yellow to reddish yellow;
capsule erect, urn cylindrical or broader
below, 2, 5-3, 2 mm long; peristome fragile,
teeth deeply cleft above short basal membrane
into two narrow, papillose processes, rarely
perforated below; operculum rostrate, 0,8
mm long; calyptra cucullate, 10 mm long;
spores round, 17-18 //m, papillose. Fig. 24:
1-9.
Ditrichum difficile is known from Asia, Micro-
nesia, India, East Africa, the Mascarenes and South-
ern Africa. In the Flora area, the species is frequently
collected in the southwestern and southern Cape and
Natal, but it has also been collected in the north-
western and eastern Cape, Zululand, Swaziland and
the eastern Transvaal. Map 32.
Vouchers: Crosby & Crosby 8167; Magill
3527; Magill & Schelpe 3923; Potts 8541; Sim 9714.
This species is somewhat variable, but only a few
collections have been difficult to place, since it is
rarely found sterile. The leaves of D. difficile are
considerably more flexuose than in any of the other
three Southern African species. In addition, the
cylindrical capsule on the very long seta, will identify
this species in Southern Africa. The capsule length
varies but does not overlap with the other species.
Ditrichum amoenum (Thwait. & Mitt.) Par. has
been excluded from the Flora. The specimens recorded
as this species in Southern Africa show a fragile,
broken peristome, not the rudimentary peristome
attributed to the Sri Lanka species. In fact, the peri-
stomes of all Southern African specimens of Ditrichum
proved quite fragile and also variable in the degree of
perforation.
Insufficiently Known Species
Ditrichum capense (C. Mull.) O. Kuntze, Rev.
Gen. PI. 2: 835 (1891). Basionym: Leptotrichum
capense C. Mull., Syn. Muse. 453 (1849). Syntypes:
Cape, Van Kamp’s Bay (Camps Bay), Ecklon s.n.;
Natal, Port Natal (Durban), Gueinzius s.n. The
syntypes have not been seen. Dixon in J. Bot., Lond.
51 : 326 (1913) refers the species to D. difficile (Dub.)
Fleisch.
Ditrichum vallis-gratiae (Lindb.) Hampe in
Flora, Jena 50: 182 (1867). Basionym: Diaphanophyl-
lum vallis-gratiae Lindb. in Ofvers. K. VetenskAkad.
Forh. 19: 605 (1863). Type: Cape, Gnadenthal,
Breutel s.n. The type has not been seen. Dixon in J.
Bot., Lond. 51: 326 (1913) refers the species to D.
difficile (Dub.) Fleisch.
94
Ditrichaceae
4. CERATODON
Ceratodon Brid., Bryol. Univ. 1: 480(1826); Broth, in Natiirl. PflFam. 10: 163 (1924); Sim,
Bryo. S. Afr. 149 (1926). Lectotype species: C. purpureus (Hedw.) Brid., vide Britt., N. Amer.
FI. 15: 60(1913).
Plants medium-sized, forming tufts; terricolous or saxicolous. Stems erect, with central
strand. Leaves lanceolate; apex acute; margins revolute, serrate at apex. Costa percurrent;
with dorsal and ventral stereid bands. Laminal cells generally quadrate, smooth.
Dioicous. Seta long; capsule cylindrical, erect to nodding, asymmetrical, strumose, fur-
rowed dry; peristome teeth deeply cleft; operculum conical; calyptra cucullate.
A widespread genus with 19 species, Ceratodon is represented in Southern Africa by C. purpureus. Species
of Ceratodon occur throughout temperate to alpine regions on all continents, including the Antarctic.
Ceratodon purpureus {Hedw.) Brid.,
Bryol. Univ. 1: 480 (1826); Broth, in Natiirl.
PflFam. 10: 163 (1924); Sim, Bryo. S. Afr.
150 (1926). Type: Europe.
Dicranum purpureus Hedw., Spec. Muse. 136
(1801).
Ceratodon stenocarpus Bruch & Schimp. ex C.
Mull., Syn. Muse. 1: 647 (1849); Broth, in Natiirl.
PflFam. 10: 163 (1924); Sim, Bryo. S. Afr. 150
(1926). Syntypes: Cape, Table Mountain, Ecklon
s.n.; Neelgheriensibus, Perrottet s.n.; Mexico,
Liebmann s.n.; Columbia, Funck & Schlim 473.
Plants medium-sized, in tufts, yellow-
green, reddish below; terricolous or saxi-
colous. Stems 5-20 mm high, yellow to
reddish, little branched, frequently tomentose
below; in section central strand present, large,
inner cortical cells large, thin-walled, outer
cortical cells in 3 rows, substereids. Leaves
incurved dry, spreading wet; ovate to lanceo-
late, 1,0- 1,5 mm; apex acute to short-
acuminate; margins narrowly revolute, entire
to apex. Costa percurrent to short-excurrent;
in section oval, guide cells 4, large, ventral
stereid band 2-3 cells thick, ventral surface
cells larger, incrassate, dorsal stereid band
3-4 cells thick, dorsal surface cells generally
substereids or only incrassate. Laminal cells
quadrate to short-rectangular, incrassate;
basal cells generally rectangular.
Dioicous. Perichaetia terminal; leaves
broad below, oblong, short-acuminate. Seta
15-25 mm long, yellow, reddish with age;
capsule erect to inclined, oblong-cylindrical,
asymmetrical, 1,0-1, 2 mm long, strongly
furrowed dry, strumose at base, yellow to
purple-red; peristome teeth 16, lanceolate,
divided to near base, divisions filiform with
weak nodes above, slightly papillose through-
out, orange below, hyaline above ; operculum
conical, 0,5 mm long; calyptra cucullate;
spores round, 10,0-12,5 /mi, smooth. Fig.
24: 10-19.
Almost cosmopolitan in distribution. In Southern
Africa, C. purpureus is common in the southwestern
and southern Cape and is infrequently collected in
Natal, Lesotho and Transvaal. The ecological tole-
rance of this species is quite wide, resulting in a good
deal of variation in plant size. Map 31.
Vouchers: Crosby & Crosby 8625; Esterhuysen
17301, 20228; Haygarth 32; Magill 4316.
Although somewhat variable, the strongly plicate,
generally inclined capsule of C. purpureus charac-
terizes the plant. Sterile plants are most easily recog-
nized by the evenly quadrate laminal cells and revo-
lute margins of the leaves.
Ceratodon stenocarpus is treated as a synonym of
C. purpureus; however, its status is still problematical.
There exist in Southern Africa, mostly in the Cape
Province, populations answering to the characters
generally ascribed to C. stenocarpus, i.e. erect, yellow
Fig. 24.— Ditrichum difficile (1-9): 1. habit, xl; 2. habit, x6; 3. stem in cross section, x 200; 4. leaves,
x40; 5. leaf in distal cross section, x370; 6. basal leaf cells at margin, x 170; 7. upper laminal cells, x 170;
8. leaf apex, xl70; 9. part of capsule mouth with peristome teeth and spores, xl25. Ceratodon purpureus
(10-19): 10. habit, xl; 11. habit, xlO; 12. stem in cross section, xl40; 13. leaves, x40; 14. leaf in cross
section, x600; 15. basal leaf cells at margin, x435; 16. upper laminal cells at margin, x435; 17. operculate
capsule, dry, x40; 18. deoperculate capsule, wet, x40; 19. part of capsule mouth with peristome teeth and
spores, x 125. (1-9, Cholnoky 1122; 10-19, Cholnoky 69).
Ditrichaceae
96
Ditrichaceae
capsule on long, yellow seta (vide Gangulee, 1971;
Grout, 1936). However, there also exist populations
intermediate in stature (from erect to inclined),
colour (from yellow to red-purple) and plication
(from smooth to deeply grooved). As no clear separa-
tion could be obtained and because no major game-
tophytic differences occur, the specimens are all
treated under C. purpureus.
5. SAELANIA
Saelania Lindb., Utkast Nat. Grupp. Eur. Bladmoss. 35 (1878); Broth, in Natiirl. PflFam.
10: 163 (1924); Sim. Bryo. S. Afr. 148 (1926). Type species: S. caesia (P. Beauv.) Lindb.
Plants small to medium, gregarious, glaucous; terricolous. Stems erect, branched, central
strand large. Leaves lanceolate, subulate to acuminate; margins serrate in acumen, plane
above, recurved below. Costa percurrent to short-excurrent, with dorsal and ventral stereid
bands. Laminal cells short-rectangular.
Autoicous. Seta long; capsule erect, elliptical; peristome teeth divided to base, densely
papillose; calyptra cucullate.
Saelania is a monotypic genus found on soil of rock crevices and in shallow caves throughout alpine and
subarctic regions of the Northern Hemisphere. In the Southern Hemisphere, it is known from similar habitats,
at high altitudes, in Southern Africa and New Zealand.
Saelania glaucescens ( Hedw .) Broth, in
Bomanss & Broth., Herb. Mus. Fenn. 2: 53
(1894); Broth, in Natiirl. PflFam. 10: 163
(1924); Sim, Bryo. S. Afr. 148 (1926). Type:
Sweden, Swartz s.n.
Trichostomum glaucescens Hedw., Spec. Muse.
112 (1801). Ditrichum glaucescens (Hedw.) Hampe in
Flora, Jena 50: 182 (1867).
Plants small, gregarious, blue-green,
covered with bluish ‘bloom’; on soil over
rock. Stems erect, 5-10 mm high, branched
above; in section with large central strand,
inner cortical cells enlarged, thin-walled,
outer cortical cells small, incrassate, in three
layers. Leaves erect-flexuose dry, patent wet;
lanceolate-subulate, 2, 5-3, 5 mm long; apex
acute; margins plane above, reflexed at mid-
leaf, irregularly serrate above. Costa excur-
rent; in section guide cells 4, large, ventral
stereid band to 3 cells thick, ventral surface
cells larger, incrassate, dorsal stereid band
large, 5-6 cells thick, dorsal surface cells
larger, incrassate. Laminal cells rectangular
to quadrate throughout, somewhat incrassate,
some upper lamina cells prorate.
Map 34. — • Saelania glaucescens
x Blindia magellanica
Fig. 25. — Saelania glaucescens (1-10): 1. habit x 1 ; 2. habit, with capsule and detached operculum, x 10;
3. stem in cross section, X 110; 4. leaves, x40; 5. leaf in cross section, x380; 6. basal leaf cells at margin,
x250; 7. laminal cells at margin (dorsal surface), X250; 8. upper laminal cells at margin, x250; 9. leaf apex,
x250; 10. part of capsule mouth with peristome teeth and spores, X 160. Distichium capillaceum (11-19): 11.
habit, I ; 12. habit, x 10; 13. stem in cross section, x 160; 14. leaf, x40; 15. leaf in proximal cross section,
X270; 16. basal leaf cells, x320; 17. laminal cells at margin, x320; 18. cells in leaf subula, x320; 19. part of
capsule mouth with peristome teeth and spores, X 160. (1-10, Magill 4584; 1 1-19, Esterhuysen 26184).
Ditrichaceae
98
Ditrichaceae
Autoicous. Perigonia terminal on short
lateral branches. Perichaetia terminal; leaves
subulate above oval base, 4-5 mm long.
Seta straight, 6-8 mm long; capsule erect,
elliptical, 1 ,0—1 , 5 mm long, weakly grooved
dry, yellow-brown; exothecial cells rounded-
rectangular, quadrate at mouth; peristome
0,25 mm high, teeth divided to base into
two linear processes, occasionally irregular,
strongly papillose, red-brown; operculum
long-rostrate, 0,5-0, 8 mm long; calyptra
cucullate; spores 17-20 pm, round, papillose.
Fig. 25: 1-10.
This widespread, high altitude species is known
from Europe, Asia, North America, New Zealand
and South Africa. In Southern Africa the species is
found in the grass-heath biome in the Drakensberg
of Natal, Lesotho and the Orange Free State. Map 34.
Vouchers: Magill 4586, 4589; Symons 8681;
Wager 86.
The plants can be identified by their bluish
‘bloom’ and erect, branching stems. In addition they
are restricted to shallow soils in small caves or reces-
ses at high altitudes.
6. DISTICHIUM
Distichium B.S.G., Bryol. Eur. 2: 153 (1846); Broth, in Naturl. PflFam. 10: 164 (1924); Sim,
Bryo. S. Afr. 149 (1926). Lectotype species: D. capillaceum (Hedw.) B.S.G., vide Grout, Moss
FI. N. Amer. 1: 38 (1936).
Plants slender, in dense tomentose tufts ; saxicolous. Stems elongate, densely tomentose
below, with central strand. Leaves distichous, oblong-subulate; base sheathing. Costa excur-
rent, with dorsal and ventral stereid band. Laminal cells subquadrate to rectangular, prorate.
Autoicous. Seta long; capsule erect, cylindrical, symmetrical; peristome teeth irregularly
cleft or perforated, lightly striate; operculum rostrate; calyptra cucullate.
The genus comprises 16 species, generally of Northern Hemisphere distribution. The single Southern
African species, D. capillaceum, is by far the most widespread species in the genus, being found on every conti-
nent. In Southern Africa all the recent collections have been made at high altitudes in the Drakensberg.
Distichium capillaceum (Hedw.) B.S.G.,
Bryol. Eur. 2: 156 (1846); Broth, in Naturl.
PflFam. 10: 164 (1924); Sim, Bryo. S. Afr.
149 (1926). Type: France.
Cynontodiuhi capillaceum Hedw., Spec. Muse. 57
(1801).
Plants medium to large, slender, forming
dense tufts, green above, brown below;
saxicolous. Stems erect, 10-30 mm high,
branching above, densely tomentose below;
in section elliptical, central strand present,
inner cortical cells in single row, large, thin-
walled, outer cortical cells in 2-3 rows,
smaller, incrassate. Leaves distichous, appres-
sed with squarrose-flexuose tips wet or dry;
oblong, abruptly long-subulate, 3-4 mm
long; apex acute, occasionally weakly tooth-
ed; base clasping stem, to 1 mm long;
margins weakly convolute below. Costa
Map 35. — • Distichium capillaceum
x Tristichium mirabile
Ditrichaceae
99
excurrent; in section flattened, guide cells 6-8,
small, incrassate, ventral stereid band small,
to 3 cells thick, ventral surface cells sub-
stereids, dorsal stereid band 4-8 cells thick,
dorsal surface cells substereids, with strongly
thickened outer walls. Upper laminal cells
rectangular, strongly prorate, at shoulders
irregular, triangular to quadrate, smooth;
basal cells long-rectangular to oblong-hexa-
gonal, linear below.
Autoicous. Perichaetia terminal; leaves
undifferentiated, base to 2 mm long. Seta
erect, 6-12 mm long; capsule erect, sym-
metrical, cylindrical, 1,0-1, 2 mm long,
yellowish; exothecial cells rectangular, qua-
drate at mouth; peristome teeth 0,2 mm
high, divided to near base or only perforated
below, lightly striate, red-yellow; operculum
rostrate; calyptra cucullate; spores round,
16-20//m, punctate. Fig. 25: 11-19.
This widespread species was treated as a doubtful
record by Sim (1926), but recent collections from the
Natal Drakensberg and Lesotho confirm its presence
in the Flora. Map 35.
Vouchers: Esterhuysen 26184; Magill 4530;
Smook 1099.
The Southern African specimens have been
collected on rock, at high altitude. The distichous
leaf arrangement and the strongly prorate cells of
the subula should easily place specimens both within
the family and the genus.
7. TRISTICHIUM
Tristichium C. Mull, in Linnaea 42: 435 (1879); Herzog in Flora, Jena 107: 322 (1914); Broth,
in Natiirl. PflFam. 10: 164 (1924). Type species: T. lorentzii C. Mull.
Plants slender, in tufts; saxicolous. Stems erect, branching; central strand present. Leaves
tristichous, rigidly appressed, linear; base sheathing. Costa percurrent, with dorsal stereid
band. Laminal cells linear, incrassate, smooth.
Autoicous. Seta short, erect to curved; capsules cleistocarpic or stegocarpic, gymnosto-
mous, oval to cylindrical ; operculum obliquely rostrate ; calyptra cucullate.
New to Africa, Tristichium was previously known only from South America. Although very similar game-
tophytically, the two South American species are separated by their cleistocarpic [ T . lorentzii C. Miill. ] or
stegocarpic [T. mirabile (C. Mull.) Herz.] capsules and different spore sizes. In Southern Africa the genus has
recently been collected on rock at high altitude in Lesotho.
Tristichium mirabile (C. Miill.) Herz. in
Flora, Jena 107: 324 (1914); Broth, in Natiirl.
PflFam. 10: 164 (1924). Type: Argentina,
Lorentz s.n.
Tristichiopsis mirabilis C. Miill. in Linnaea 43 : 394
(1882).
Plants medium-sized, slender, in tufts,
light green to yellow-green; saxicolous.
Stems 10-15 mm high, branching above,
weakly radiculose below; in section sub-
triangular, central strand small, inner cortical
cells in 2-3 rows, varying in size, thin-walled,
outer cortical cells in 2 rows, stereids, dark
red. Leaves tristichous, rigidly appressed wet
or dry, small and distant below, larger and
crowded above ; linear to narrowly triangular,
0,8- 1,2 mm long, ventral surface broadly
channelled to keeled; apex acute; base
sheathing stem; margins erect, entire. Costa
percurrent, superficial cells linear, smooth;
in section ventrally concave, guide cells
12-16, exposed ventrally, incrassate, smooth,
dorsal stereid band l(-2) cells thick across
costa, dorsal surface cells slightly larger,
strongly incrassate, smooth. Upper laminal
cells linear, incrassate, smooth; basal cells
similar, weakly thickened, reddish yellow at
insertion.
Gonioautoicous. Perigonia gemmate,
axillary on upper stem; leaves oval-acumi-
nate, 0,6-0, 7 mm long. Perichaetia terminal;
leaves convolute, oblong-subulate, 1,2 mm
100
Ditrichaceae
long; costa filling subula. Seta erect to
somewhat curved dry, 4 mm long, yellow to
brownish; capsules erect to curved, ±
asymmetrical, 1 ,5 mm long, smooth, brown-
ish; exothecial cells thickened; stomata not
seen; annulus revoluble; peristome absent;
operculum obliquely rostrate, 0,5 mm high,
cells not twisted; calyptra cucullate, small;
spores round, 20-25 //m, coarsely papillose,
brownish. Fig. 26: 1-14.
The species has only recently been discovered in
the moist alpine to subalpine grasslands of Lesotho.
Map 35.
Vouchers: Van Zanten 7609943a, 7609957,
7609984.
Although occasionally inconspicuous because of
branching or development of perichaetia, the dis-
tinctly tristichous arrangement of the closely appres-
sed, linear leaves will separate this species from all
other Southern African taxa. The character is most
easily observed on sterile, unbranched stems.
Fig. 26. — Tristichium mirabile (1-14): 1. habit, x 1 ; 2. habit, x 5; 3. stem in cross section, x320; 4. leaves,
x65; 5. leaf in proximal cross section, x640; 6. leaf in distal cross section, x640; 7. cells at leaf base (right
side), xl70;8. lower laminal cells, x 640; 9. upper lamina! cells, X 640; 10. leaf apex, X 170; 11. perigonium,
X65; 12. perichaetial leaf, x65; 13. exothecial cells, X640; 14. spores, X400. Blindia magellanica (15-24):
15. habit, xl; 16. habit, x5; 17. stem in cross section, x320; 18. leaves, x25; 19. leaf in proximal cross
section, ' 640; 20. leaf in distal cross section, x640; 21. basal leaf cells at right margin, x 170; 22. leaf apex,
xl70; 23. upper laminal cells, X640; 24. basal laminal cells, x640. (1-14, Van Zanten 7609884; 15-24, Van
Zanten 76091012).
Ditrichaceae / Seligeriaceae
101
103
SELIGERIACEAE
Plants very small to medium, gregarious or loosely caespitose ; saxicolous. Stems simple or
branching; central strand present. Leaves erect to falcate-secund ; generally subulate above
broader base; margins entire to serrulate. Costa percurrent to excurrent, occasionally filling
subula; in section cells not strongly differentiated. Laminal cells rectangular to linear, smooth;
basal cells generally longer; alar cells not distinct or strongly differentiated.
Perichaetial leaves somewhat larger. Seta erect to cygneous; capsule small, ovoid to
ellipsoid or pyriform, often with broad mouth; peristome single; operculum rostrate;
calyptra cucullate ; spores essentially smooth.
A small family of seven genera, of which the largest, Blindia, is the only one known from Southern Africa.
BLINDIA
Blindia B.S.G., Bryol. Eur. 2: 17 (1846); Broth, in Nattirl. PflFam. 10: 170 (1924); Sainsb.,
N. Zeal. Moss. 87 (1955); Smith, Moss FI. Brit. Irel. 125 (1978). Type species: B. acuta (Hedw.)
B.S.G.
Plants of medium size, yellowish green to brownish green, frequently dark brown to
blackish below, glossy ; generally saxicolous and semi-aquatic. Stem slender, simple or branched ;
central strand present. Leaves crowded, patent to falcate-secund; subulate above oblong,
concave base. Costa narrow below, filling subula; in section cells not differentiated. Laminal
cells rectangular to oblong-hexagonal, occasionally sinuolate ; alar cells strongly differentiated,
coloured.
Dioicous or autoicous. Perichaetial leaves occasionally distinct. Seta long or short, erect
and flexuose to cygneous; capsule exserted or immersed, short-pyriform to subspherical ;
peristome teeth short-lanceolate, smooth, rarely absent; operculum obliquely beaked; calyptra
cucullate ; spores large, 20-32 p m.
A genus of 33 species, Blindia is most widely distributed in the Southern Hemisphere, especially in colder
regions.
Blindia magellanica Schimp. ex C. Mull.
in Bot. Ztg 20: 328 (1862); Broth, in Nattirl.
PflFam. 10: 171 (1924); Sainsb., N. Zeal.
Moss. 87 (1955); Van Zanten in Van Zinderen
Bakker et al. [eds], Marion and Prince
Edward Islands 184 (1971). Type: South
America, Tierre del Fuego, Insula Eremitae,
Hooker s.n.
Plants of medium size, loosely caespi-
tose, glossy, yellow-green above, brownish
below; semi-aquatic. Stems 10-15 mm high,
occasionally branched; in section subround,
central strand very small, inner cortical cells
in 4 rows, incrassate, outer cortical cells
smaller, in 2 rows, stereids. Leaves crowded,
weakly secund above, erect below wet or
dry; lanceolate-subulate, 2, 5-4, 5 mm long,
concave below; apex acute; margins entire,
erect to incurved. Costa filling subula,
superficial cells linear; in section subelliptical,
flattened ventrally, cells weakly differentiated
proximally, homogeneous in subula. Upper
laminal cells incrassate, short- to long-
rectangular, somewhat sinuolate, smooth or
dorsal surface rough (obvious only in cross
section); cells in subula short-rectangular;
basal cells long-rectangular to oblong-
hexagonal, sinuolate; alar cells strongly
differentiated, quadrate, rectangular at mar-
gins, incrassate, reddish.
Sporophytes not known from Southern
Africa. The generally smaller Marion Island
plants have: perichaetial leaves undifferen-
tiated; seta flexuose dry, cygneous wet, 4
104
Seligeriaceae
mm long; capsule short-elliptical, 0,8 mm
long, mouth broad; peristome teeth 16, short-
triangular, smooth; operculum obliquely
rostrate, 0,6 mm high; spores round, 20 fim.
Fig. 26: 15-24.
New to Southern Africa, B. magellanica is
known from southern South America, Australia,
New Zealand and the subantarctic islands. In South-
ern Africa the species has only recently been collected
on rock, at 3200 m, in northern Lesotho. Map 34.
Voucher: Van Zanten 76091012.
The species is most easily recognized by its
strongly differentiated alar cells, the absence of
specialized marginal leaf cells and by the cells of the
costa in cross section being practically homogeneous.
The plants and leaves of the Southern African speci-
men are larger than the Marion Island plants.
105
DICRANACEAE
Plants minute to large, gregarious or in dense cushions; terricolous or saxicolous. Stems
erect, frequently tomentose below; central strand present or absent. Leaves appressed to
spreading or squarrose, rarely falcate, secund; generally narrow, elongate, rarely ligulate or
very short-oval, or consisting of modified multi-layered costa; bases occasionally clasping;
margins plane or convolute, occasionally reflexed above, entire to dentate, sometimes bistra-
tose; frequently bordered by several rows of narrow, elongate, hyaline cells. Costa strong or
occasionally very weak, frequently very broad below, ending below apex to filling subula, or
excurrent as long, denticulate awn; in section round to oblong, guide cells generally differen-
tiated, ventral stereid band present or absent, ventral cells sometimes enlarged, dorsal stereid
band generally strong or stereids in small groups separated by larger incrassate cells, dorsal
surface smooth to rough, mammillose or with lamellae 1-6 cells long, or costa strongly
modified, consisting of single layer of small, rhombic chlorocysts, surrounded by 2-8 layers of
large, thin-walled leucocysts. Upper laminal cells quadrate to rectangular, occasionally elon-
gate, incrassate, smooth or papillose, rarely mammillose or prorate, infrequently pitted; basal
cells moderately differentiated, large, rectangular; alar cells occasionally differentiated,
enlarged, reddish.
Perichaetia terminal or lateral through innovation ; leaves rarely differentiated, occasion-
ally base long-sheathing. Seta very short or elongate; capsule stegocarpous, peristomate or
gymnostomous, or rarely cleistocarpous; urn short-elliptical to long-cylindrical, frequently
ribbed, neck occasionally differentiated, long or short; peristome when present single, rudi-
mentary to well developed, teeth generally cleft to middle, vertically striate or barred below,
papillose above; operculum short- or long-rostrate; calyptra cucullate, base entire or rarely
fringed; spores mostly small, infrequently highly ornate.
A large family comprising 64 genera, of which 14 are known from Southern Africa.
Key to Subfamilies of Dicranaceae
1 Alar cells conspicuous, enlarged, hyaline or coloured.. .Subfamily Dicranoideae (p. 126)
1 Alar cells not differentiated :
2 Leaf cells strongly papillose, mammillose or prorate:
3 Leaves squarrose above clasping base Subfamily Dicranoideae (p. 126)
3 Leaves erect to spreading, not squarrose :
4 Leaf cells prorate Subfamily Trematodontoideae (p. 106)
4 Leaf cells papillose or mammillose Subfamily Dicranoideae (p. 126)
2 Leaf cells essentially smooth :
5 Leaves in section with small triangular or rhombic chlorocysts surrounded by
enlarged, hyaline leucocysts Subfamily Leucobryoideae (p. 156)
5 Leaf cells not differentiated into chlorocysts and leucocysts :
6 Stems julaceous, leaves appressed when wet or dry :
7 Leaf margins entire; costa weak, consisting of only a few undifferentiated,
thin-walled cells Subfamily Trematodontoideae (p. 106)
7 Leaf margins erose-dentate; costa stronger, consisting of 3 rows of stereids or
substereids Subfamily Dicranelloideae (p. 118)
106
Dicranaceae
6 Leaves patent with flexuose tips when dry, spreading when wet:
8 Costa very broad, occupying | to \ width of leaf base
Subfamily Dicranoideae (p. 126)
8 Costa narrower, occupying less than ^ width of leaf base :
9 Costa excurrent as short, distinct awn; capsules pendent on flexuose seta
Subfamily Dicranoideae (p. 126)
9 Costa not excurrent as distinct awn, frequently percurrent or filling subula ;
capsule erect, seta weakly flexuose :
10 Capsules short-elliptical, without differentiated neck, generally ribbed
when dry; leaves mostly abruptly subulate above oval or obovate base
or stems elongate, leaves distant, ovate, acuminate to subulate
Subfamily Dicranelloideae (p. 118)
10 Capsules elliptical to cylindrical with well defined neck, occasionally
longer than urn ; leaves acuminate to gradually subulate
Subfamily Trematodontoideae (p. 106)
Subfamily Trematodontoideae
Plants minute to medium, gregarious or in loose tufts; terricolous. Stems erect; central
strand present. Leaves patent, ovate to oblong, acuminate to subulate or appressed, oval with
broadly rounded apices; margins entire to serrate above. Costa strong or weak, narrow,
flattened; in section with or without stereid or substereid bands. Laminal cells quadrate to
rectangular, smooth or prorate; basal cells rectangular; alar cells not differentiated.
Perichaetia terminal; leaves erect, occasionally distinct from vegetative leaves. Capsule
stegocarpic or cleistocarpic, immersed to exserted, with well developed neck; peristome absent,
rudimentary or well developed; teeth with vertical bars below, perforated or cleft to below
middle; operculum rostrate ; calyptra cucullate or mitriform; spores large, papillose to spinose.
The genera treated in this subfamily would be included in the family Bruchiaceae, recently restored by
Buck (1979).
Key to Genera of Subfamily Trematodontoideae
1 Plants small to medium ; costa in section with guide cells and stereid or substereid bands ;
capsule exserted, neck long, narrow 3. Trematodon
1 Plants minute to small; costa in section without internal differentiation; capsule im-
mersed, neck short, broad, with numerous large stomata:
2 Leaves erect, apex acuminate; capsule cleistocarpic 1. Bruchia
2 Leaves appressed, apex round; capsule stegocarpic 2. Cladophascum
1. BRUCHIA
Bruchia Schwaegr., Spec. Muse. Suppl. 2: 91 (1824); Broth, in Naturl. PflFam. 10: 173 (1924);
Sim, Bryo. S. Afr. 151 (1926). Type species: B. vogesiaca Schwaegr.
Plants very small to minute, gregarious or in small groups; terricolous. Stems very short;
central strand present. Leaves patent; ovate-acuminate, generally bistratose above. Costa
percurrent to short-excurrent; in section cells weakly differentiated. Laminal cells quadrate to
rectangular, smooth.
Fig. 27. — Cladophascum gymnomitrioides (1-9): 1. habit, X 1 ; 2. habit, x 20; 3. stem in cross section, x 150;
4. leaves, x 50; 5. leaf in cross section, x 435; 6. leaf showing cells, x 85; 7. perichaetial leaves, x 30; 8. sporo-
phyte with spores and calyptra, x 60; 9. spores, x 260. Bruchia brevipes (10-20): 10. habit, xl; 11. habit, x20;
12. stem in cross section, x60; 13. leaf, x60; 14. leaf in cross section, x435; 15. basal leaf cells at margin,
xl70; 16. leaf apex, xl70; 17. perichaetial leaf, x43; 18. capsule with spores, x20; 19. calyptra, x20; 20.
spore, x 426. (1-2 & 7-9, Volk 882; 3-6, Sim 8747; 10-20, Mag ill & Schelpe 4017).
Dicranaceae
107
108
Dicranaceae
Autoicous or dioicous. Perigonia gemmate, axillary or terminal on minute plants; peri-
chaetia terminal; leaves oval-subulate to oblong-acuminate. Capsule cleistocarpic, immersed
to emergent; urn pyriform or elliptical, neck well developed; calyptra mitriform; spores round
to reniform, spinose or punctate.
The 24 species of Bruchia are scattered through the temperate regions of both hemispheres. By far the
greatest number of species, namely 12, is known from North America, followed by southern South America
with 5 species. Three species are known from Southern Africa.
1 Spores 40-45 /jm; spinose 1. B. brevipes
1 Spores 25-33 pm; not spinose:
2 Spores 30-33 pm; distal surface pitted, proximal surface with strong tetrad scar 2. B.foveolata
2 Spores 25-30 ^m; papillose to vermiculate, tetrad scar weak 3. B. eckloniana
1. Bruchia brevipes Harv. ex Hook, in
Hooker’s, Icon. PI. 3: 231 (1840); Broth, in
Natiirl. PflFam. 10: 173 (1924); Sim, Bryo.
S. Afr. 152 (1926). Type: Cape, Newlands,
Harvey s.n. (BM, holo. !; MANCH!).
Phascum elegans Hornsch. in Linnaea 15: 114
(1841). Bruchia elegans (Hornsch.) C. Miill. in Bot.
Ztg 5: 99 (1847); Roth in Hedwigia 53: 92 (1913).
Type: Cape, Table Mountain, Ecklon s.n. (H-BR!).
Plants minute, gregarious, light green;
terricolous. Stems erect, to 0,5 mm high ; in
section with small central strand, cortical
cells in ± 3 rows, uniform in size, incrassate,
reddish. Leaves erect, bistratose above base;
oval to ovate, acuminate to subulate in sub-
perichaetial leaves, 1-2 mm long; margins
plane, entire to serrulate by projecting cells.
Costa excurrent, awn of upper leaves serru-
late; ventral and dorsal superficial cells
elongate, frequently prorate; in section guide
cells 4, central, ventral cells in one layer,
large, dorsal cells in 2 layers, large, incras-
sate. Laminal cells narrowly rectangular
above shoulders; basal cells broader, rec-
tangular to oblong-hexagonal.
Autoicous. Perigonia axillary, leaves
oval-acuminate, 0,8 mm long. Perichaetial
leaves oval-subulate, 2,5 mm long. Seta
erect, 0, 3-0,4 mm long; capsule cleisto-
carpic, pyriform, to 1 mm long, neck with
numerous stomata, to 0,5 mm long; calyptra
mitriform, covering upper urn; spores round-
ed to subreniform, spinose, 40-45 pm,
yellowish. Fig. 27: 10-20.
Endemic to the fynbos biome of the southern
and southwestern Cape, specimens of B. brevipes are
rarely collected, perhaps because of their small size
and ephemeral nature. The original collection was
made in Cape Town; recent collections have been
made in the mountains of the Clanwilliam area.
Map 36.
Map 36. — • Bruchia brevipes
x Bruchia foveolata
Vouchers: Magill & Schelpe 4017; Schelpe 7710.
The plants grow on crusts of soil in association
with Pleuridium and Fissidens. The small plants and
cleistocarpic, pyriform capsules are distinctive and
are diagnostic for the genus. Vegetatively the three
Southern African species are very similar, although
B. brevipes is slightly larger than the other two. The
spores offer the only reliable characters for separating
the three species.
2. Bruchia foveolata Magill, sp. nov., B.
queenslandica Stone habitu sculpturaque spo-
rarum similis, sed statu dioico, marginibus
foliar um reflexis irregulariter serratis et
sporis majoribus differt.
Type: South West Africa/Namibia,
Windhoek, Farm Rietfontein, on soil, Volk
01160o (PRE, holo.; herb. Volk; MO).
Plants very small, scattered, yellow-
green; terricolous. Stems 0,2-0, 6 mm tall,
simple; in section subround, central strand
present, cortical cells in 3-4 rows, outer cells
Dicranaceae
109
weakly thickened. Leaves larger above,
iappressed dry, erect-spreading wet; lanceo-
late to linear-lanceolate, subulate, 0,5-1, 5
mm long; margins reflexed, irregularly ser-
rate by projecting cells, bistratose above.
Costa short-excurrent; ventral and dorsal
superficial cells rectangular, smooth; in
section elliptical, guide cells not differentiated
or 2 lateral cells larger, ventral and dorsal
cells round, incrassate. Upper laminal cells
rectangular to triangular, incrassate, smooth;
basal cells weakly differentiated, rectangular,
thin-walled, hyaline.
Dioicous. Male plants smaller, perigonia
terminal. Perichaetia terminal; leaves larger,
oval-subulate, 2,0-2, 5 mm long. Seta 0,1-
0,2 mm long, hyaline; capsule cleistocarpic,
pyriform-rostrate, 1 mm long, yellowish,
Fig. 28.— Bruchia foveolata (1-12): 1. habit, xl; 2. habit, x26; 3. stem in cross section, x210; 4. leaf,
x60; 5. leaf in proximal cross section, x435; 6. leaf in distal cross section, x435; 7. basal leaf cells at margin,
xl70; 8. upper laminal cells, x440 ; 9. leaf apex, xl70; 10. exothecial cells and stomata of capsule neck,
x200; 11. calyptra, x26; 12. spores, x640. B. eckloniana (13-22): 13. habit, xl; 14. habit, x40; 15. leaves,
x80; 16. leaf in distal cross section, x640; 17. leaf base, x 170; 18. upper laminal cells, x200; 19. leaf apex,
X 1 70; 20. exothecial cells and stomata of capsule neck, x200; 21. calyptra, x40; 22. spore, x640. (1-12,
Volk 01160a; 13-22, Ecklon s.n.).
110
Dicranaceae
beak to 0,2 mm long, neck short, 0,3 mm
long, stomata superficial (phaneropore); exo-
thecial cells quadrate, thin-walled; calyptra
mitriform, lobed below, smooth, 0,6 mm
long, covering urn ; spores subround to reni-
form, 30-33 pm, distal surface pitted, proxi-
mal surface with strong tetrad scar, bright
light yellow. Fig. 28: 1-12.
This new species was collected on sandy soil in
the Khomas Highland Savanna of central South
West Africa/Namibia. Map 36.
Voucher: Type only.
In addition to the pitted spores with strong
tetrad scars, B. foveolata differs from the other
Southern African species by a large calyptra that
completely covers the urn and the reflexed, irregularly
serrate leaf margins.
3. Bruchia eckloniana C. Mull., Syn.
Muse. 1 : 19 (1848); Broth, in Natiirl. PfIFam.
10: 173 (1924); Sim, Bryo. S. Afr. 152 (1926).
Type: Cape, Ecklon s.n. (H-BR!).
Plants very small, scattered or in small
groups, yellow-green; terricolous. Stems 0,2-
0,4 mm high, simple; in section subround,
central strand present, cortical cells in 3-4
rows. Leaves crowded, larger above, erect-
spreading wet or dry; ovate-acuminate,
0, 5-1,0 mm long, bistratose above; margins
plane, entire. Costa percurrent to short-
excurrent; ventral and dorsal superficial
cells elongate, smooth; in section flattened,
guide cells 2-4, weakly differentiated, ventral
cells incrassate, in 1-2 rows, dorsal cells
subquadrate, in 3 rows. Upper laminal cells
quadrate to rectangular, incrassate, smooth;
basal cells rectangular, incrassate, hyaline.
Dioicous. Perigonia terminal. Perichae-
tia terminal; leaves oblong-acuminate, 1,2-
1,5 mm long. Seta 0,2 mm long, hyaline;
capsule cleistocarpic, cylindrical-rostrate, 1
mm long, yellow, beak to 0,2 mm long, neck
short, 0,2 mm long, stomata superficial
(phaneropore) ; exothecial cells quadrate,
thin-walled; calyptra mitrate, 0,4 mm long,
covering upper capsule only; spores sub-
round to reniform, 25-30 pm, vermiculate to
papillose, red-brown. Fig. 28: 13-22.
The exact type locality is not given by Muller
(1849) who cites only ‘Prom. bon. spei: Ecklon’.
Since additional collections have not been seen,
the exact distribution of this species is not known.
Map 37.
Voucher: Type only.
The spores of B. eckloniana are papillose or
vermiculate when the papillae are united into irregular
ridges. The small calyptra and the plane, entire leaf
margins will help to identify the species.
2. CLADOPHASCUM
Cladophascum Dix. ex Sim, Bryo. S. Afr. 143 (1926). Type species: C. gymnomitrioides (Dix.)
Dix. ex Sim.
Plants small, gregarious or in loose patches; terrestrial. Stem erect, sparsely branched;
central strand present. Leaves appressed; oval, apex rounded. Costa weak, short; in section
cells undifferentiated. Laminal cells quadrate to angular.
Perichaetia conspicuous; capsule immersed, stegocarpous ; peristome absent; operculum
conical; calyptra small, mitriform; spores large, papillose.
The monotypic genus Cladophascum is restricted to the central and western parts of Southern Africa and
southern Zimbabwe.
Cladophascum gymnomitrioides (Dix.)
Dix. ex Sim, Bryo. S. Afr. 143 (1926). Type:
Zimbabwe, Matopos, Sim 8850 (BM, lecto. !,
selected here; PRE!).
Aongstroemia gymnomitrioides Dix. in S. Afr. J.
Sci. 18: 301 (1922); Broth, in Natiirl. PflFam. 10:
179 (1924).
Plants small, in loose tufts, yellow-green;
terricolous. Stems julaceous, erect, to 10 mm
high, irregularly branched; in section round,
central strand present, small, inner cortical
cells large, subround, thin-walled, outer
cortical cells quadrate to rectangular, in 1-2
rows, incrassate. Leaves imbricate, concave,
semicircular to oval, 0,2-0, 4 mm long;
apex broadly obtuse to rounded ; base
clasping; margins erect, entire. Costa weak to
above mid-leaf; in section consisting of 4
undifferentiated cells. Upper laminal cells
smooth, quadrate to angular, strongly incras-
sate; upper marginal cells short-rectangular,
thin-walled, hyaline, forming border 2-3 cells
Dicranaceae
111
wide around apex; basal cells larger, quadrate
to transversely rectangular, hyaline.
Synoicous. Perigonial leaves small, oval,
obtuse. Perichaetia large, lateral through
innovation; leaves broadly oval to oblong,
abruptly cuspidate; costa strong above,
excurrent, weak to absent in lower base. Seta
very short, c. 0,1 mm long; capsule stego-
carpous, pyriform, 0,6 mm long, neck short,
0,2 mm long, with numerous stomata; peris-
tome absent; operculum conical, very short-
beaked; calyptra mitriform, covering oper-
culum; spores subreniform, 52-57 pm,
lightly papillose. Fig. 27 : 1-9.
These small plants are rarely collected. They
grow on soil in grassland and savanna in southern
Zimbabwe, southern Transvaal, Orange Free State,
northern Cape and South West Africa/Namibia.
Map 37.
Vouchers: Sim 9706, 9707; Van Rooy 485;
Volk 882, 6094; Wager B155.
Sterile plants are characterized by the sparsely
branched, julaceous stem. The large-leaved, lateral
perichaetia of fertile plants are very distinctive.
Gametophytically the plants are very distinct from
any other genus in the family, however, the capsule
morphology clearly indicates the correct placing of
Map 37. — • Cladophascum gymnomitrioides
x Bruchia eckloniana
Cladophascum in Trematodontoideae with Bruchia
and Trematodon.
The lectotype ( Sim 8850) was originally erro-
neously numbered 8450. The mistake was corrected
in a letter from Sim to Dixon (PRE) and it was
published as 8850. The error was, however, not cor-
rected on the specimens.
3. TREMATODON
Trematodon Michx. in FI. Bor. Am. 2: 289 (1803); Broth, in Natiirl. PflFam. 10: 174 (1924);
Grout, Moss FI. N. Amer. 1: 37 (1936); Sim, Bryo. S. Afr. 152 (1926). Type species: T.
longicollis Michx.
Plants very small to medium, gregarious or in loose tufts; terricolous or saxicolous.
Stems erect, simple; central strand present. Leaves subulate above oblong to oval base;
margins frequently bistratose. Costa generally subpercurrent, with or without ventral stereid
band. Leaf cells rectangular, smooth or prorate.
Perichaetia terminal; leaves oval to oblong, subulate. Capsule exserted; urn cylindrical,
neck as long as or longer than urn, frequently strumose; operculum long rostrate; calyptra
cucullate.
Trematodon is a genus of c. 82 species found mostly in the Southern Hemisphere. The genus has been
divided into two subgenera (cf. Brotherus, 1924), namely Gymnotrematodon with peristome absent or only a
basal membrane present, and Trematodon with peristome well developed. Six species are known from Southern
Africa, equally divided between the two subgenera.
Emphasis has been placed on sporophyte characters by most authors (Bartram, 1939; Gangulee, 1971),
as is the case here. This should not cause problems, since specimens of Trematodon, in Southern Africa, are
rarely collected without capsules.
1 Peristome absent or rudimentary (Subgenus Gymnotrematodon) :
2 Leaf cells prorate 1. T. pillansii
2 Leaf cells smooth :
3 Upper leaves subulate; apex acute; cells of leaf margin narrowly rectangular, 7-10 pm wide; peri-
stome rudimentary 2. T. intermedius
3 Upper leaves lingulate; apex obtuse; cells of leaf margin bulging rectangular, 15-18 pm wide;
peristome absent 3. T. paradoxus
112
Dicranaceae
1 Peristome well developed (Subgenus Trematodon ):
4 Leaves lanceolate to ligulate, obtuse 6. T. mayottensis
4 Leaves oblong to oval, abruptly subulate:
5 Capsule neck ± as long as urn; spores with distinct tetrad scar, 30-35 pm 4. T. divaricatus
5 Capsule neck longer than urn; spores without distinct scar, 22-25 pm 5. T. longicollis
1. Trematodon pillansii Dix. ex Sim,
Bryo. S. Afr. 153 (1926). Type: Cape,
Miller’s Point, Pillans 4058 (BM, lecto.!,
selected here; BOL!; PRE!).
Plants very small, gregarious, yellowish
green; saxicolous or terricolous. Stems to
1 mm high; in section central strand very
small, cortical cells incrassate, slightly smaller
toward margin, reddish. Leaves spreading
wet and dry; oblong to oval, subulate, 1,0-
1,5 mm long; apex obtuse; margins crenate,
dentate at apex; lamina bistratose above.
Costa ending below apex; in section guide
cells 4, central, ventral cells in 1-2 layers,
strongly incrassate, dorsal cells in 2-3 layers,
substereids or strongly incrassate. Lamina l
cells rectangular, occasionally quadrate at
shoulders, proximal and distal ends prorate;
basal cells larger, rectangular, smooth.
Map 38. — • Trematodon paradoxus
X Trematodon pillansii
Perichaetial leaves oval, abruptly ligu-
late, 2,5 mm long; base sheathing. Seta
erect, 4-5 mm long, yellow; capsule erect,
yellowish, urn 1,0- 1,2 mm long, neck 1,5-
2,0 mm long, without stroma, exothecial
cells short, hexagonal; peristome absent;
operculum rostrate, 0,5 mm high; calyptra
covering upper urn; spores round, 70-75 p m,
proximal face with tetrad ridge, distal face
with C-shaped papillae. Fig. 29: 1-9.
Described from the Cape Peninsula, T. pillansii
is endemic to the southwestern Cape. The species
grows on shallow soil and bare rock in the fynbos
biome. Map 38.
Vouchers: Barnard 50333; Esterhuysen 24363;
Magill & Schelpe 4068; Pillans 4060; Sim 9282,
9297.
The small size of these plants, prorate leaf cells,
absence of peristome and the large spores are diag-
nostic for this species. These characters also indicate
that T. pillansii is not closely related to the other
Southern African species.
2. Trematodon intermedius Welw. &
Duty in Mem. Soc. Phys. Hist. nat. Geneve
21: 226 (1870); Broth, in Natlirl. PflFam.
10: 175 (1924); Dix. & Wag. in Trans. R.
Soc. S. Afr. 18: 248 (1930). Type: Angola,
Huilla, Lopollo River, Welwitsch 9 (G,
holo. !).
Plants small, gregarious, green; terri-
colous. Stems 1-2 mm tall; in section central
strand small, inner cortical cells lax, outer
cortical cells small, incrassate, reddish.
Leaves flexuose to erect wet or dry; oval to
oblong, subulate, 1, 5-3,0 mm long; apex
acute, bluntly toothed; margins entire, bistra-
tose above. Costa percurrent to short-
excurrent; in section elliptical, guide cells 4,
central, exposed laterally, ventral stereid
Fig. 29. — Trematodon pillansii (1-9): 1. habit, xl;2. habit, X 10; 3. stem in cross section, x 230; 4. leaves,
x30; 5. leaf in cross section, x550; 6. basal leaf cells, x435; 7. upper laminal cells at margin, x435; 8. leaf
apex, x 435; 9. part of capsule mouth and spores, x 35. T. intermedius (10-18): 10. habit, xl; 11. habit, xlO;
12. leaves, x25; 13. leaf in cross section, x275; 14. basal leaf cells at costa, x200; 15. upper laminal cells at
margin, x200; 16. leaf apex, x200; 17. part of capsule mouth with rudimentary peristome and spores, x35;
18. operculum, X 10. T. paradoxus (19-27): 19. habit, xl; 20. habit, X 10; 21. stem in cross section, x230;
22. leaves, : 25; 23. leaf in cross section, x 500; 24. basal leaf cells at costa, x400; 25. upper laminal cells at
margin, 400; 26. leaf apex, x400; 27. part of capsule mouth and spores, x35. (1-9, Magill 4068; 10-11, Sim
8187; 12-18, Kemp 746; 19-27, Sim 7213).
Dicranaceae
113
114
Dicranaceae
band small, of 6-8 cells in central group,
ventral surface cells small, incrassate, only
over stereid band, dorsal stereid band in 1-2
rows across costa, dorsal surface cells larger,
completely covering stereid band. Laminal
cells rectangular above shoulders ; basal cells
lax, rectangular to oblong-hexagonal.
Perichaetial leaves oblong-subulate, 2,0-
2,5 mm long. Seta erect, 3-4 mm long,
yellow ; capsule erect, yellow, urn short-cylin-
drical, to 1 mm long, neck to 1,5 mm long,
strumose; peristome rudimentary, consisting
of short basal membrane, occasionally with
short, irregular, papillose thickenings; oper-
culum rostrate, to 1 mm long; calyptra
cucullate, small, not completely covering
urn; spores rounded, 25-28 n m, papillose.
Fig. 29: 10-18.
Originally described from Angola, T. intermedius
is also known from east Africa south into Zimbabwe,
the central and eastern Transvaal, Swaziland, eastern
Orange Free State, Natal, Zululand, southern Cape
and northern South West Africa/Namibia. Map 39.
Map 39. — • Trematodon intermedius
x Trematodon divaricatus
Vouchers: Cholnoky 242; Kemp 746; Rehmann
439; Sim 8450: Wager 184; Volk 2225; Welwitsch
20.
Sim’s (1926) treatment of T. intermedius as a
synonym of T. paradoxus is incorrect. The leaves of
the two species, especially the shape of the subula
and upper leaf cells, are quite distinct, as noted by
Brotherus (1924) and Dixon & Wager (1930). In
addition, the peristome is completely absent in T.
paradoxus but present, although rudimentary, in T.
intermedius.
3. Trematodon paradoxus Hornsch. in
Linnaea 15: 122 (1841); Broth, in Natiirl.
PflFam. 10: 176 (1924); Dix. & Wag. in
Trans. R. Soc. S. Afr. 18: 248 (1930).
Type: Cape, Table Mountain, Ecklon s.n.
(BM!).
Plants very small, in scattered groups,
green; terricolous. Stems 1-2 mm high; in
section central strand small, inner cortical
cells large, thin-walled, outer cortical cells
in 2 rows, small, incrassate, red. Leaves erect,
weakly flexuose dry, erect-spreading wet;
oval to oblong, abruptly linear, 1, 5-2,0 mm
long; apex obtuse, occasionally acute; mar-
gins plane, entire, unistratose. Costa sub-
percurrent, in section guide cells 4, exposed
ventrally, dorsal stereid band 1-2 cells thick,
dorsal surface cells larger, weakly thickened,
occasionally extending upward interrupting
stereid band. Laminal cells bulging, rectangu-
lar above shoulders, in 1-3 rows on either
side of costa at apex; marginal cells bulging;
basal cell long-rectangular or oblong-hexa-
gonal, lax.
Perichaetial leaves oblong-subulate, 2,0-
2,5 mm long. Seta erect to flexuose dry, 2-3
mm long, yellow; capsule erect, yellow, urn 1
mm long, neck 0,8- 1,0 mm long, strumose;
peristome absent; operculum rostrate, 0,6-
0,8 mm long; calyptra cucullate, covering
operculum and capsule mouth; spores ±
round, 30-35 nm, coarsely papillose. Fig.
29: 19-27.
Known from the Central African Republic,
Zimbabwe and South Africa, Trematodon paradoxus
was described from Table Mountain. Additional
material from the southwestern Cape has not been
seen. Recent specimens have been collected in the
southeastern Transvaal, eastern Orange Free State,
Zululand, Natal, and the eastern Cape. Map 38
Vouchers: Cholnoky 251; Owen 27; Sim 9725,
196; Smook 1098; Tolken 5739.
This species resembles T. divaricatus in many
respects; however, T. paradoxus is generally smaller
and lacks a peristome. The spores of this species
also approach those of T. divaricatus. but they are
not as highly ornamented and the tetrad scar is weak,
when present.
4. Trematodon divaricatus Bruch ex
Krauss in Flora, Jena 29: 133 (1846); Broth,
in Natiirl. PflFam. 10: 176 (1924). Type:
Natal, Umgeni (Mgeni) River, Krauss s.n.
(BM!).
Dicranaceae
115
Trematodon aequicollis Ren. & Card, in Bull. Soc.
r. Bot. Belg. 41 : 10 (1905); Broth, in Natiirl. PflFam.
10: 176 (1924); Sim, Bryo. S. Afr. 156 (1926). Type:
Zaire, Kisantu, Gillet s.n. (PC, holo.!).
Plants small, in loose tufts, green; terri-
colous. Stems 3-4 mm high, simple; in
section central strand small, inner cortical
cells large, thin-walled, outer cortical cells in
1-2 rows, smaller, incrassate, reddish. Leaves
contorted dry, erect-flexuose wet; oblong to
elliptical, abruptly subulate, 2,0-2, 5 mm
long; apex obtuse to acute; margins plane,
bluntly toothed at apex. Costa broad below,
ending below apex; in section flattened, guide
cells exposed ventrally or with 2-3 small,
ventral surface cells, dorsal stereid or sub-
stereid band 2 cells thick, dorsal surface cells
smaller than guide cells, surface irregular,
some cells bulging outward. Upper lamina l
cells short- rectangular to quadrate; marginal
cells at shoulders and lower subula quadrate ;
basal cells large, lax, rectangular or oblong-
hexagonal.
Perichaetial leaves similar to vegetative
leaves, base broadly oval, clasping, 2, 5-3,0
mm long. Seta erect, 10-13 mm long, yellow;
capsule erect to nodding, brown, urn cylin-
drical, 2, 5-3,0 mm long, neck to 1,5 mm
long, weakly strumose; peristome reddish,
teeth triangular, 0,4-0, 5 mm high, perforated
below, cleft into two filaments above, with
vertical bars below, papillose apically, hyaline
and papillose fringe between teeth forming
ornate border on filaments and lower parts
of teeth; operculum rostrate, to 1 mm high;
calyptra cucullate, covering capsule; spores
round, 30-35 pm, spinose, with distinct
tetrad scar. Fig. 30: 1-11.
Trematodon divaricatus is known from eastern,
western and Southern Africa. In the Flora area, it has
been collected in the northeastern Transvaal but is
more common from Zululand southwards through
Natal to Port St Johns in the Transkei. Map 39.
Vouchers: Van der Bijl 27; Lambert 17; Schelpe
7522; Sim 9736; Wager 1474; Wood 7127.
This species is recognized by its wide leaf cells,
short-necked capsule, peristome divided into fila-
ments above and its strongly spinose spores with
distinct tetrad scar. It can be separated from the
very similar but smaller T paradoxus by longer leaves,
broader costa and larger, peristomate capsule.
5. Trematodon longicollis Michx. in FI.
Bor. Am. 2: 289 (1803); Broth, in Natiirl.
PflFam. 10: 176 (1924); Bartram in Philipp.
J. Sci. 68: 28 (1939); Gangulee, Moss. E.
India 2: 231 (1971). Type: North America.
Trematodon pallidens C. Mull, in Linnaea 40:
242 (1876); Broth, in Natiirl. PflFam. 10: 176 (1924);
Sim, Bryo. S. Afr. 155 (1926). Type: Comoro Islands,
Hildebrandt 1812 (BM!).
Trematodon flexifolius C. Mull, in Flora, Jena 69:
278 (1886); Broth, in Natiirl. PflFam. 10: 176
(1924); Sim, Bryo. S. Afr. 155 (1926). Type: Sao
Tome, Henriquez s.n.
Trematodon africanus Wag. & Dix. in Trans. R.
Soc. S. Afr. 4: 4 (1914); Sim, Bryo. S. Afr. 154 (1926).
Type: Transvaal, Tzaneen, Wager s.n., PRE-CH12159
(PRE, lecto.!, selected here; BM!).
Plants small to medium, gregarious or in
loose tufts, green; terricolous. Stems 1-7 mm
high; in section central strand large, cortical
cells large, lax, becoming smaller with thicker
walls toward margin. Leaves contorted above
erect base dry, erect to spreading wet; oval
to oblong, abruptly subulate, 2-4 mm long;
apex bluntly acute; base sheathing; margin
bistratose in subula, with a few projecting
cells above, apex frequently toothed. Costa
subpercurrent to percurrent; in section
flattened, guide cells 6, ventral stereid band
small, 3-4 cells in single central row, ventral
surface cells smaller than guide cells, covering
entire costal surface or restricted to area
above ventral stereid band and exposing
guide cells laterally, dorsal stereid band 1-3
cells thick, extending across costa, dorsal
surface cells smaller than guide cells, occa-
sionally some cells larger, extending upward
Map 40. — • Trematodon longicollis
x Trematodon mayottensis
116
Dicranaceae
Dicranaceae
117
interrupting stereid band. Laminal cells short-
rectangular to quadrate above shoulders;
basal cells large, lax, rectangular to oblong-
hexagonal.
Perichaetial leaves oblong-subulate, 3-4
mm long. Seta 6-15 mm long, yellowish;
capsule erect, yellow, urn cylindical, 1 ,2-1 ,6
mm long, neck 2, 2-3, 5 mm long, strumose;
peristome reddish, teeth lanceolate, 340-360
fim high, perforated, without free filaments,
vertically barred below, upper segments
papillose, yellowish, fringe present between
teeth, weak, papillose, hyaline; operculum
long-beaked, 1,0-1, 5 mm high; calyptra
cucullate; spores round, 22-25 n m, bluntly
papillose. Fig. 30: 21-29.
Cosmopolitan; in Southern Africa, this species
has been collected in the northern, central and south-
ern Transvaal, eastern Orange Free State and Natal.
Map 40
Vouchers: Bottomley 1338; Brenan M2785; Sim
8559, 9730; Wager 92, 273, 1107.
The wider definition of this species employed by
Bartram (1939) and Gangulee (1971) is adopted here.
A capsule neck that is consistently twice as long as
the urn, strongly perforated peristome teeth without
free filaments, and papillose spores without obvious
tetrad scar, separate T. longicollis from the other
species. Although gametophytically similar to several
other species, identification has not been a problem,
as all collections examined have numerous sporo-
phytes.
6. Trematodon mayottensis Besch. in
Annls Sci. nat. Bot., ser. 7, 2: 84 (1885);
Broth, in Natiirl. PflFam. 10: 176 (1924);
Sim, Bryo. S. Afr. 154 (1926). Type: Mayotte
Island, Marie 1 (BM, holo. !).
Trematodon ligulatus Rehm. ex Roth in Hedwigia
53: 96 (1913). Type: Natal, Oakford, Rehmann 22
(PRE!).
Plants small, gregarious or in loose
tufts, light green; terricolous. Stems (1— )2— 3
mm high, simple; in section central strand
large, cortical cells large, becoming incrassate
toward margin. Leaves contorted dry, erect-
spreading wet; lanceolate to ovate-ligulate,
1,0-1, 6 mm long; apex obtuse to rounded;
margins weakly reflexed, entire to crenulate
above, bistratose; in section marginal cells
rounded, smaller than rectangular laminal
cells. Costa ending below apex to subper-
current; in section flattened, cells not strongly
differentiated, guide cells somewhat larger,
ventral cells in single layer, thin-walled, dorsal
substereid band 1-2 cells thick, cells variable
in size and thickening occasionally only
incrassate, dorsal surface cells similar to
ventral cells. Laminal cells smooth, short-
rectangular to quadrate, thin-walled; basal
cells larger, rectangular, lax.
Perichaetial leaves not differentiated.
Seta erect, 3 mm long, yellow-brown;
capsule erect, yellow-brown, urn short-
cylindrical, 1, 8-2,0 mm long, neck 1 mm
long, strumose at base; peristome reddish
yellow, teeth triangular, to 240 //m high,
irregularly perforated, with vertical bars
below, apical segments free, papillose, hya-
line; operculum long-beaked, 0,6 mm high;
calyptra cucullate; spores subreniform, 22-27
Hm, papillose. Fig. 30: 12-20.
In Southern Africa T. mayottensis is known from
Oakford in Natal and Wolhuter Kop in the Orange
Free State. The type is from Mayotte Island; addi-
tional specimens have been collected in Zimbabwe.
Map 40.
Voucher: Wager 92.
Dixon & Wager (1930) disagreed with Sim’s
(1926) placing of T. ligulatus Rehm. ex Roth under T.
mayottensis. Examination of the specimens involved,
showed that the specimens of Rehmann 22, in South
African herbaria, are T. mayottensis, while the speci-
men at BM is T. longicollis (Dixon in Sim corresp.
in PRE as T. africanus). Trematodon mayottensis is
separated from other species of Trematodon in
Southern Africa, by its broad leaves and obtuse
apices.
Fig. 30. — Trematodon divaricatus (1-11): 1. habit, xl; 2. habit, xlO; 3. stem in cross section, x 1 35 ;
4. leaves, x 35; 5. leaf, x 50; 6. leaf in cross section, x435; 7. basal leaf cells at costa, x435; 8. upper laminal
cells at margin, x240; 9. leaf apex, x435; 10. part of capsule mouth with peristome teeth and spores, x35;
11. spores, x 300. T. mayottensis (12-20): 12. habit, x 1 ; 13. habit, x 10; 14. stem in cross section, x 135; 15.
leaves, x 35; 16. leaf in cross section, x435; 17. basal leaf cells at costa, x220; 18. upper laminal cells at costa,
x220; 19. leaf apex, x220; 20. part of capsule mouth with peristome teeth and spores, x35. T. longicollis
(21-29): 21. habit, xl; 22. habit, xlO; 23. stem in cross section, xl35; 24. leaves, x35; 25. leaf in cross
section, x 435; 26. basal leaf cells at costa, x 220; 27. upper laminal cells at margin, x 220; 28. leaf apex, x220;
29. part of capsule mouth with peristome teeth and spores, X35. (1-11, Van der Bijl 27; 12-20, Rehmann 22;
21-29, Wager 1107c).
118
Dicranaceae
Subfamily Dicranelloideae
Plants small, caespitose, yellow-green; terricolous, or saxicolous. Stems erect, slender,
occasionally julaceous; central strand present. Leaves appressed, spreading or squarrose;
apex acuminate to subulate or rounded; margins plane, entire to erose-denticulate. Costa
filling subula, percurrent or ending below apex, with dorsal stereid band. Laminal cells rec-
tangular to rhomboidal, occasionally quadrate, smooth ; alar cells not differentiated.
Dioicous or rarely monoicous. Perichaetia terminal, leaves not distinct or bases elongate.
Seta erect; capsule short, elliptical to cylindrical, frequently ribbed dry; peristome present or
absent, teeth cleft to middle, striate below or irregularly perforated, papillose throughout;
operculum rostrate; calyptra cucullate.
Key to Genera of Subfamily Dicranelloideae
1 Stems julaceous; leaves appressed wet or dry 1. Aongstroemia
1 Stems not julaceous; leaves flexuose or occasionally appressed dry, erect-spreading wet:
2 Leaves abruptly subulate above broad base, 1 ,8-6,0 mm long 2. Dicranella
2 Leaves lanceolate, gradually acuminate, 1,0-1, 8 mm long:
3 Plants small; leaves closely set; stems smooth 3. Microdus
3 Plants long, slender; leaves distant; stems papillose 2. Dicranella
1. AONGSTROEMIA
Aongstroemia B.S.G., Bryol. Eur. 1: 171 (1846); Broth, in Natiirl. PflFam. 10: 179 (1924);
Sim, Bryo. S. Afr. 157 (1926). Type species: A. longipes (Somm.) B.S.G.
Plants small to medium, caespitose, yellow-green; terricolous. Stems slender, julaceous;
central strand present. Leaves appressed wet or dry, oval to oblong; apices rounded or subu-
late; margins plane, entire to erose-denticulate. Costa ending below apex or filling subula; in
section cells undifferentiated or guide cells exposed ventrally, dorsal stereid band well de-
veloped. Laminal cells elongate, incrassate.
Dioicous. Perichaetia terminal. Setae erect; capsules ovoid-cylindric ; peristome present or
absent; operculum rostrate.
The genus contains c. 15 species of temperate or alpine distribution. Two species occur in Southern Africa
and are recognized by the small, slender, julaceous stems with closely appressed leaves wet or dry.
1 Leaves oval; apices rounded, apiculate; margins erose-denticulate; costa ending below apex. . . .2. A.julacea
1 Leaves oblong; apices subulate; margins entire; costa extending into subula 1. A. filiformis
1. Aongstroemia filiformis ( P . Beauv.)
Wijk & Marg. in Taxon 9: 50 (1960). Type:
Madagascar.
Dicranum filiforme P. Beauv., Prodr. 53 (1805).
Weissia vulcanica Brid., Muscol. Recent. Suppl.
1: 124 (1806). Aongstroemia vulcanica (Brid.) C.
Mull., Syn. Muse. 1: 427 (1848); Broth, in Natiirl.
PflFam. 10: 179 (1924); Sim, Bryo. S. Afr. 157 (1926).
Type: Bourbon.
Aongstroemia transvaaliensis C. Miill. in Hedwigia
38: 89 (1899); Broth, in Natiirl. PflFam. 10: 179
(1924). Type: Transvaal, Spitzkop, Wilms s.n. (G,
holo.!).
Plants small to medium, loosely caespi-
tose, yellow-green; terricolous. Stems jula-
ceous, 10-20 mm high; in section subround,
central strand of medium size, inner cortical
cells in 2-3 rows, thin-walled, outer corti-
cal cells in 2-4 rows, stereids. Leaves appres-
sed with flexuose subulae wet or dry ; oblong,
1,4-1, 8 mm long; apex gradually subulate;
base sheathing; margins plane, entire. Costa
filling subula; in section guide cells 6-8,
exposed ventrally, dorsal stereid band strong,
in 3 rows, dorsal surface cells substereids,
Dicranaceae
119
Fig. 31. — Aongstroemia filiformis (1-9): 1.
habit, xl; 2. habit, x5; 3. stem in cross section,
x290; 4. lower stem leaves, x40; 5. upper stem
leaves, x40; 6. leaf in cross section, xl20; 7. basal
leaf cells at margin, Xl70; 8. leaf apex, xl70; 9.
upper laminal cells, x640. A. julacea (10—19) : 10.
habit, xl; 11. habit, x5; 12. stem in cross section,
x290; 13. leaves, x40; 14. leaf in cross section,
x250; 15. basal leaf cells at margin, x 170; 16. basal
leaf cells, x640; 17. upper laminal cells, x640; 18.
upper leaf margin, x435; leaf apex, xl70; 19. leaf
apex, x 170. (1-9 Cholnoky 465; 10-19, Sim 9738).
120
Dicranaceae
with minute, blunt papillae covering dorsal
surface of costa and lamina. Upper laminal
cells rhomboidal to fusiform, weakly sinuo-
late, strongly incrassate, minutely papillose
dorsally (visible in section only); basal cells
rhomboidal to rectangular, weakly thickened.
Sporophyte not known in Southern
Africa. Fig. 31 : 1-9.
Found in East Africa, Madagascar and Reunion ;
in Southern Africa A. filiformis is rarely collected in
forests of the eastern Transvaal and Natal. Map 41.
Voucher: Cholnoky 465.
The tall, slender plants and leaves with appressed
bases and flexuose subulae are characteristic for this
species.
2. Aongstroesnia julacea (Hook.) Mitt, in
J. Linn. Soc., Bot. 12: 27 (1869); Broth, in
Natiirl. PflFam. 10: 179 (1924); Sim, Bryo.
S. Afr. 157 (1926). Type: Central America,
Andes, Humboldt & Bonpland s.n.
Gymnostomum julaceum Hook., Musci Exot. 1 : 42
(1918).
Plants small, caespitose, green to yellow-
green; terricolous. Stems slender, julaceous,
5-10 mm high, simple; in section round,
central strand of medium size, inner cortical
cells large, in 3 rows, weakly thickened, outer
cortical cells in 1-2 rows, stereids, reddish.
Leaves appressed wet or dry; short-oval,
0,4-0, 8 mm long; apex rounded-apiculate to
acute; margins plane, erose-denticulate by
projecting cell ends of adjoining marginal
cells. Costa ending below apex; in section
cells undifferentiated, in 3 rows, stereids and
substereids. Upper laminal cells rectangular
Map 41. — • Aongstroemia filiformis
x Aongstroemia julacea
to rhombic, incrassate; basal cells slightly
larger, quadrate, weakly thickened.
Sporophyte not known in Southern
Africa. Fig. 31: 10-19.
Aongstroemia julacea is known from Central
America, northern South America, Southern Africa,
the East African Islands, India, northern Asia and
Japan. In Southern Africa the species is only rarely
collected in alpine grassland in the Drakensberg of
Natal and Lesotho. Map 41.
Vouchers: Edwards PRE-CH4600; Sim 8686.
Although smaller, the julaceous stems of A.
julacea macroscopically resemble Anomobryum. The
leaf shape, costal development and anatomy, and
leaf cell shape separate the genera. This species is
recognized by its oval leaves with erose-denticulate
margins and the costa ending below the apex.
2. DICRANELLA
Dicranella (C. Mull.) Schimp., Coroll. 13 (1856); Broth, in Natiirl. PflFam. 10: 181 (1924);
Sim, Bryo. S. Afr. 158 (1926). Lectotype species: D. grevilleana (Brid.) Schimp., vide Grout,
Moss FI . N. Amer. 1 :54 (1936).
Aongstroemia sect. Dicranella C. Mull., Syn. Muse. 1 : 430 (1848).
Plants small, scattered or in dense tufts, yellowish green; terricolous or saxicolous.
Stems simple or occasionally branched; central strand present. Leaves patent to squarrose;
acuminate or abruptly subulate above oval or oblong to obovate, clasping base; margins plane,
entire. Costa percurrent; ventral stereid band present or absent. Laminal cells quadrate to
rectangular, incrassate, smooth; basal cells rectangular, thin-walled; alar cells not differen-
tiated.
Dicranaceae
121
Dioicous or rarely monoicous. Perichaetia terminal, leaves similar to vegetative leaves,
bases longer sheathing. Seta long, erect, straight, yellow to red-brown with age; capsule short-
cylindrical, erect to curved, slightly ribbed dry; peristome erect, teeth cleft to middle, verti-
cally striate below, red-yellow, papillose and hyaline above; operculum long-rostrate; calyptra
cucullate; spores round, papillose.
The genus Dicranella contains approximately 60 species, found throughout temperate and tropical regions.
The genus is infrequently collected in the eastern and southern parts of Southern Africa.
1 Leaves (2-) 4-6 mm long, subulate above oblong clasping base, lamina unistratose at mid-leaf
1 . D. subsubulata
1 Leaves 0,8-2, 2 mm long; lamina bistratose at mid-leaf:
2 Leaves patent, ovate to lanceolate, lamina irregularly bistratose above base 3. D. rigida
2 Leaves squarrose above oblong, clasping base, regularly bistratose above base 2. D. symonsii
1. Dicranella subsubulata (C. Mull.)
Jaeg. in Verh. St Gall, naturw. Ges. 1870-71 :
375 (1872); Broth, in Natiirl. PflFam. 10:
182 (1924); Sim, Bryo. S. Afr. 159 (1926).
Type: Cape, Gnadenthal, Ecklon s.n. (BM,
holo. !).
Aongstroemia subsubulata Hampe ex C. Miill. in
Bot. Ztg 16: 162 (1858).
Campylopus nanus C. Miill. in Bot. Ztg 5: 804
(1857). Dicranum nanum (C. Miill.) C. Miill., Syn.
Muse. 1: 383 (1849). Microcampylopus nanus (C.
Miill.) Broth, in Natiirl. PflFam. 10: 183 (1924);
Dixon in Trans. R. Soc S. Afr. 8: 185 (1920); Sim,
Bryo. S. Afr. 180 (1926), non Dicranella nana Wint. in
Hedwigia 55: 85 (1914). Type: Cape, Gnadenthal,
Drege s.n. (BM !).
Aongstroemia borgeniana Hampe in Bot. Ztg 28:
33 (1870). Dicranella borgeniana (Hampe) Jaeg. ex
Par., Ind. Bryol 326 (1896); Broth, in Natiirl. PflFam.
10: 182 (1924). Type: Natal, Mapumulo, Borgen s.n.,
1867 (BM, holo.!; HBG!).
Aongstroemia abruptifolia C. Miill. in Hedwigia 38:
89 (1899). Dicranella abruptifolia (C. Miill.) Par., Ind.
Bryol. 2: 6 (1903); Broth, in Natiirl. PflFam. 1 : 310
(1909).Type: Cape, Esternek, Rehmann 25 (BOL!;
NH!; PRE!).
Microcampylopus pusillus C. Miill. in Hedwigia 38:
78 (1899); Dix. in Trans. R. Soc. S. Afr. 8: 185
(1920); Sim, Bryo. S. Afr. 180 (1926). Type: Cape,
Montagu & Outeniqua Pass, Breutel s.n. (BM !).
Anisothecium natalense P. Varde in Revue bryol.
lichen. 24: 29 (1955). Syntypes: Natal, Royal Natal
National Park, Cholnoky 67, 11 (PC!).
Plants small, loosely caespitose, green to
yellow-green; terricolous. Stems 2-4(-8) mm
long, simple; in section round, central strand
present, inner cortical cells in 3-4 rows,
thin-walled, outer cortical cells in 2 rows,
smaller, incrassate. Leaves flexuose above
base when dry, erect-spreading wet; abruptly
subulate above broad, oblong to obovate or
occasionally oval base, (2-)4-6 mm long;
apex weakly toothed; base sheathing; margins
plane, entire; in section lamina unistratose.
Costa filling subula, dorsal and ventral
superficial cells rectangular, smooth; in
proximal section concave, guide cells 8,
exposed laterally, ventral stereid band small,
ventral surface cells small, incrassate, dorsal
stereid band 4-5 cells thick, dorsal surface
cells substereids, smooth; in distal section
ventral stereid band small, ventral surface
cells incrassate, dorsal stereid band 2-3
cells thick, occasionally some cells not
stereids, dorsal surface cells incrassate,
smooth. Laminal cells rectangular, incras-
sate, smooth; basal cells rectangular to
rhomboidal, thin-walled.
Perichaetia terminal, leaves to 6 mm
long, sheathing bases elongate. Seta erect,
to 5 mm long, yellowish to red-yellow;
capsules short-cylindrical, 1,5 mm long,
weakly plicate dry, red-brown; exothecial
cells rectangular, incrassate, quadrate to
hexagonal at mouth; peristome erect, teeth
triangular, 0,3 mm long, cleft to middle,
vertically striate below, red-yellow, papillose
and hyaline above; operculum long-rostrate,
1 , 1 mm long, cells not twisted; spores round,
30 /rm, papillose, yellow-brown. Fig. 32: 1-12.
This species is presently known from eastern and
Southern Africa. In the Flora area, D. subsubulata is
infrequently collected in grassland and forest road
cuttings or soil banks in the southwestern, southern
and eastern Cape, Transkei, Natal, Lesotho, eastern
Orange Free State, Zululand and the eastern and
northern Transvaal. Map 42.
Vouchers: Cholnoky 390, 487, 904; Magill 4111.
The leaves of D. subsubulata are generally
abruptly subulate above a broad, sheathing base.
Leaves of smaller plants are occasionally more
gradually constricted to the subula and have oval,
rather than the typical oblong to obovate bases. The
smooth, erect to flexuose subula will help to dis-
tinguish this species from D. symonsii.
122
Dicranaceae
Map 42. — • Dicranella subsubulata
2. Dicranella symonsii Dix. in Trans.
R. Soc. S. Afr. 8: 183 (1920); Sim, Bryo. S.
Afr. 160 (1926). Type: Natal, Giant’s Castle,
Symons sub Sim 8665 (PRE !).
Anisothecium symonsii (Dix.) Broth, in Natiirl.
PflFam. 10: 177 (1924).
Plants slender, forming cushions, green
to yellow-green ; saxicolous. Stems 20-40 mm
high, infrequently branched, with dense
reddish tomentum below; in section sub-
round, central strand present, inner cortical
cells in 3-4 rows, large, thin-walled, outer
row smaller, incrassate. Leaves squarrose
above clasping base wet or dry, flexuose
above when dry; abruptly long-acuminate
above oblong or obovate, clasping base,
1,8-2, 2 mm long; apex weakly toothed;
base sheathing stem; margins plane, entire;
in section lamina bistratose, ventral cells
larger. Costa not well defined, percurrent, to
| width of leaf base, ventral superficial
cells long-rectangular, smooth, dorsal super-
ficial cells rectangular, mammillose; in
distal section guide cells 6, large, ventral
stereid band weak, in single layer, ventral
surface cells small, rounded, incrassate,
smooth, dorsal stereid band weak, in single
layer or small groups separated by guide
cells, dorsal surface cells vertically elongate,
projecting dorsally. Laminal cells rectangular
to short-rectangular, slightly thickened, papil-
lose with low, weak papillae at both ends of
cells in acumen; basal cells moderately
differentiated, rectangular, somewhat larger,
weakly thickened.
Sporophyte unknown. Fig. 32: 13-20.
Endemic to Southern Africa, D. symonsii is
rarely collected on rock in the montane grasslands of
the Drakensberg in Lesotho, Orange Free State and
Natal. Map 43.
Voucher: Esterhuysen 20213.
The long stems, sheathing leaf bases and squar-
rose, papillose acumens separate D. symonsii from
the other species. The plants could perhaps be mis-
taken for a species of Bartramia, especially in view of
the weak papillae at both ends of the cells in the
acumen (prorate). The size of the plants, habit and
anatomy seem to indicate the correct placing of these
specimens in Dicranella, although sporophytes are
needed before a final assessment can be made.
3. Dicranella rigida Dix. ex Sim, Bryo. S.
Afr. 160 (1926). Type: Cape, Paarl, Sim
9633 (BM, holo. !; PRE!).
Map 43. — • Dicranella rigida
X Dicranella symonsii
Fig. 32. — Dicranella subsubulata (1-12): 1. habit, xl; 2. habit, xlO; 3. stem in cross section, x260; 4.
leaves, x 40 ; 5. leaf in proximal cross section, x 435 ; 6. leaf in distal cross section, x 435 ; 7. cells at leaf shoulder,
x 170; 8. laminal cells at shoulder, x 640; 9. upper laminal cells, x 640; 10. leaf apex, x 170; 11. part of capsule
mouth with peristome teeth, xl20; 12. spore, x550. D. symonsii (13-20): 13. habit, xl; 14. habit, x5; 15.
stem in cross section, x260; 16. leaves, X40; 17. leaf in distal cross section, x435; 18. cells at leaf shoulder,
x 170; 19. leaf apex, x 170; 20. upper laminal cells, x640. D. rigida (21-27): 21. habit, x 1 ; 22. habit, x5;
23. stem in cross section, x260; 24. leaves, x40; 25. leaf in median cross section, x435; 26. upper laminal
cells, X 640; 27. leaf apex, x 170. (1-12, Cholnoky 962; 13-20, Sim 866; 21-27, Sim 9633).
Dicranaceae
123
124
Dicranaceae
Plants slender, forming loose mats,
green to yellow-green; terricolous. Stems
10-30 mm long, ascending, rarely branched,
papillose; in section round, central strand
small, inner cortical cells in 3-4 rows,
large, outer cortical cells in 2 rows, small,
incrassate, cells of outer row with a single
sharp papilla. Leaves distant, erect-spreading
to flexuose dry, erect-spreading wet; ovate to
lanceolate or triangular, short-acuminate to
subulate, 0,8-1, 8 mm long; base not sheath-
ing; margins plane, entire; in section lamina
bistratose in patches between costa and
margin, cells of similar size, dorsal cells
weakly bulging. Costa percurrent, ventral
superficial cells long-rectangular, smooth,
dorsal superficial cells rectangular, rough;
in section guide cells 6, ventral cells in 2 rows,
incrassate, dorsal cells in 2-3 rows, incras-
sate, dorsal surface cells slightly elongate
vertically, bulging. Laminal cells quadrate,
rectangular or triangular, thin-walled, smooth
or mammillose, 3-4 rows of cells at insertion
slightly larger, rectangular to rhomboidal.
Sporophyte unknown. Fig. 32: 21-27.
Endemic to Southern Africa, D. rigida is rarely
collected in the fynbos biome of the southwestern
Cape. Map 43.
Vouchers: Brenan M2763; Garside 6461, 6622.
The long, slender stems and small, distant leaves
give the plants an etiolated appearance. The leaves
vary in size and shape in the few specimens seen.
The habit, papillose stem, rectangular, thin-
walled leaf cells and leaf anatomy give the plants a
distinctive appearance, but D. rigida remains a
troublesome species to identify.
3. MICRODUS
Microdus Schimp. ex Besch. in Mem. Soc. Sci. nat. Cherbourg 16: 161 (1872); Broth, in
Natiirl. PflFam. 10: 181 (1924); Gangulee, Moss. E. India 247 (1971). Type species: not
designated.
Plants small, tufted; terricolous. Stems with central strand. Leaves appressed to erect-
spreading, narrow; lanceolate to elliptical, acuminate; margins plane, entire. Costa percurrent,
with dorsal stereid band. Upper laminal cells quadrate to rhomboidal, weakly thickened ; alar
cells not differentiated.
Dioicous. Perichaetia terminal, leaves not distinct. Seta erect; capsule short, oval to
cylindrical; peristome teeth somewhat irregular, cleft or perforated, papillose; operculum long-
rostrate; spores small, papillose.
A genus of c. 60 species found primarily in tropical regions throughout the world. The genus is rare in
temperate regions, as is the case in Southern Africa.
Microdus minutus ( Hampe ) Besch. in
Annls Sci. nat. Bot., ser. 6, 9: 305 (1880);
Broth, in Natiirl. PflFam. 10: 181 (1924).
Type: Madagascar, Borgen sub 22 (BM,
holo. !).
Aongstroemia minuta Hampe in Linnaea 38: 209
(1874). Dicranella minuta (Hampe) Jaeg. in Verh. St
Gall, naturw. Ges. 1877-78: 375 (1880); Sim, Bryo. S.
Afr. 158 (1926).
Plants small, loosely caespitose, green to
yellow-green; terricolous. Stems 2-4 mm high,
rarely branched; in section round, central
strand small, inner cortical cells in 2-4 rows,
thin-walled, outer row weakly thickened,
reddish. Leaves weakly appressed dry, erect-
spreading wet; lanceolate or oblong to ellip-
tical, acuminate, 1 ,0-1,4 mm long; margins
plane, entire; in section lamina infrequently
with small, bistratose patches. Costa per-
current; in section crescent-shaped, guide
cells 6, ventral cells absent or 1-2 large,
thin-walled cells over central guide cells,
dorsal stereid or substereid band 1-2 cells
thick, dorsal surface cells incrassate, weakly
bulging, occasionally all cells except on
dorsal surface undifferentiated, strongly in-
crassate. Upper laminal cells quadrate, rect-
angular or rhomboidal, weakly thickened,
smooth; basal cells moderately differentiated,
rectangular, thin-walled or slightly thickened.
Perichaetia terminal; leaves slightly lar-
ger. Seta erect, to 4 mm long, red-brown;
capsules elliptical, 0,6-0, 8 mm long, con-
stricted below mouth when dry, red-brown to
yellow-brown, exothecial cells rectangular.
Dicranaceae
125
Fig. 33. — Microcampylopus perpusillus (1-8): 1. habit, X 1; 2. habit, X 10; 3. stem in cross section, X350;
4. leaves, x40; 5. leaf in cross section, x 170; 6. basal leaf cells, x 170; 7. leaf apex, x 170; 8. upper laminal
cells at margin, x 640. Microdus minutus (9-16): 9. habit, x 1 ; 10. habit, x 10; 11. stem in cross section, x 350;
12. leaves, x40; 13. leaf in cross section, x435; 14. basal leaf cells, xl70; 15. upper laminal cells at margin,
X640; 16. leaf apex, x 170. (1-8, Sim 8789; 9-16, Magill 3421).
126
Dicranaceae
Map 44. — • Microdus minutus
x Microcampylopus perpusillus
weakly thickened; peristome irregular, yel-
low-brown, fragile, teeth cleft and perforated,
50-150 high, ornately papillose through-
out; operculum rostrate, 0,8 mm long, cells
not twisted ; calyptra cucullate ; spores round,
20 nm, yellow-brown, papillose. Fig. 33:
9-16.
This species is presently known from the East
African Islands, and from eastern and Southern
Africa. In the Flora area it is infrequently collected in
grasslands of the central and southern Transvaal,
Swaziland and Natal. The species has also been col-
lected in the southern and southwestern Cape. Map
44.
Vouchers: Cholnoky 248; Magill 3421, 4279.
This species could easily be mistaken for a
Dicranella, but it is distinguished by short, narrow,
lanceolate to elliptical leaves without pronounced
shouldeis, by non-ribbed, oval capsules, and rudi-
mentary peristomes that are papillose throughout.
Subfamily Dicranoideae
Plants small to large, in loose tufts, dark green, yellow-green or light green; terricolous,
saxicolous or corticolous. Stems erect, occasionally tomentose below; central strand present
or absent. Leaves erect to falcate-secund, generally narrow, elongate; lamina unistratose;
margins serrate to entire, frequently bordered by several rows of narrow, hyaline cells or
marginal cells undifferentiated. Costa percurrent to excurrent, narrow or occupying up to f
of leaf base; in section with dorsal and ventral stereid bands or ventral cells thin-walled,
frequently strongly differentiated. Laminal cells quadrate, rectangular, rhomboidal or rarely
linear, smooth, mammillose or papillose, occasionally pitted; basal cells rectangular; alar
cells frequently strongly differentiated.
Perichaetia terminal or lateral through innovation, leaves undifferentiated or base long-
sheathing. Seta long, erect, flexuose or curved; capsules cylindrical; peristome teeth well
developed, cleft to middle, vertically striolate below, or infrequently irregularly perforated to
rudimentary; operculum rostrate; calyptra cucullate, base entire or fimbriate.
Key to Genera of Subfamily Dicranoideae
1 Alar cells not differentiated:
2 Costa broad, flattened in cross section:
3 Ventral cells of costa numerous, large, completely covering guide cells .... 6. Campylopus
3 Ventral cells of costa few, small, covering only central guide cells. . . 5. Microcampylopus
2 Costa narrow, rounded in cross section:
•4 Costa ending below apex to percurrent; margins irregularly serrate; leaf cells
sharply mammillose dorsally and ventrally 1 . Oreoweisia
4 Costa mucronate; margins entire; alar cells distinct but usually broken away; leaf
cells domed ventrally, smooth dorsally or weakly papillose juxtacostally
2. Holomitrium
Dicranaceae
127
1 Alar cells distinct:
5 Leaves with differentiated border of narrow, hyaline cells; costa narrow:
6 Upper leaf cells strongly papillose, with large stellate or spinose papillae or with
numerous small papillae scattered over lumen; leaf cells generally quadrate
above base 4. Leucoloma
6 Upper leaf cells smooth, mostly elongate 3. Dicranoloma
5 Leaves without distinct hyaline border:
7 Laminal cells domed ventrally, smooth dorsally or juxtacostal cells weakly papil-
lose; costa narrow 2. Holomitrium
7 Laminal cells smooth dorsally and ventrally; costa very broad, generally with lamel-
lae or sharply mammillose cells on dorsal surface:
8 Leaves falcate-secund ; dorsal costal cells sharply mammillose; basal laminal cells
only weakly differentiated 7. Chorisodontium
8 Leaves rigidly erect; dorsal costa smooth, rough or with lamellae, cells not
sharply mammillose; basal laminal cells moderately differentiated. . . 6. Campylopus
1. OREOWEISIA
Oreoweisia ( B.S.G .) De Not. in Epil. 1 : 489 (1869); Broth, in Natiirl. PflFam. 10: 197 (1924);
Sim, Bryo. S. Afr. 250 (1926). Type species: O. serrulata (Funck) De Not.
Weissia subgenus Oreoweisia B.S.G. , Bryol. Eur. 1 : 71 (1846).
Plants small, in tufts; terricolous. Stems erect, occasionally branched; central strand
present. Leaves strap-shaped ; apex obtuse ; margins bluntly serrate. Costa pronounced, ending
just below apex; in section round, with dorsal stereid band only. Laminal cells irregularly
quadrate, mammillose, incrassate.
Oreoweisia is a widespread genus with 18 species. The highest concentration of species (8) is found in
South America. Two species are known from Africa, O. bruntonii (Sm.) Mild, of northern Africa and Europe
and the Southern African endemic O. erosa. The latter was treated by Sim (1926) under Pottiaceae; careful
examination, however, reveals its relationship to Dicranaceae. In the absence of sporophytes, the erose leaf
margins, strongly mammillose leaf cells and costal anatomy will help to identify specimens.
Oreoweisia erosa (C. Mm//.) Kindb.,
Enum. Bryin. Exot. 69 (1888); Broth, in
Natiirl. PflFam. 10: 198 (1924); Sim, Bryo.
S. Afr. 250 (1926). Type: Cape, Ecklon s.n.
Weissia erosa Hampe ex C. Miill. in Bot. Ztg 16:
163 (1858).
Plants small, in loose tufts, dark green
to yellow-green; terricolous. Stems erect, to
10 mm tall, radiculose below; in section
subround, central strand present, inner
cortical cells in 3 rows, large, outer cortical
cells in 1-2 rows, small, incrassate, reddish.
Leaves crisped when dry, patent wet; ligulate,
1, 2-2,0 mm long; apex obtuse, occasionally
acute, frequently weakly cucullate; margins
plain, erose to bluntly serrate above. Costa
prominent, ending just below apex or rarely
percurrent; in section guide cells 4, large,
ventral cells in single row, incrassate, dorsal
stereid band 2 cells thick, dorsal surface
cells larger, incrassate. Laminal cells irregular,
quadrate to angular, sharply mammillose on
dorsal and ventral surfaces; basal cells
oblong, smooth, hyaline; alar cells not
differentiated.
Autoicous. Perigonia on lower stem,
leaves abruptly ligulate from a broadly oval
base. Perichaetia lateral through innovation,
frequently persistent; leaves not differen-
tiated. Seta erect, 5-6 mm long, reddish
yellow; capsule short-cylindrical, 1,5 mm
long, reddish yellow; peristome fragile,
Dicranaceae
Dicranaceae
129
teeth irregularly perforated, yellowish,
smooth; spores round, 20-22 pm, papillose.
Fig. 34: 1-9.
Endemic to Southern Africa. Originally described
from the southwestern Cape, O. erosa has been
infrequently collected in scattered locations in Natal,
Lesotho and Transvaal. Only a single collection was
seen from the southwestern Cape. Map 45.
Vouchers: Esterhuysen 15365; Magill 4307,
4486; Sim 8047, 10059; Wager 1441.
Despite the obvious problem of a macroscopic
resemblance to taxa of Pottiaceae, it is unlikely that
O. erosa will be confused with any other pottiaceous or
dicranaceous species. The distinctly erose upper leaf
margins and the strong mammillae on both dorsal and
ventral cell surfaces will help to identify the species.
Dixon (in Sim, 1926) compares the type of O. erosa
to the Northern Hemisphere species O. torquescens
(Brid.) Wijk & Marg. ( = 0 . serrulata (Funck)
De Not.).
2. HOLOMITRIUM
Holomitrium Brid., Bryol. Univ. 1 : 226 (1826); Broth, in Natiirl. PflFam. 10: 201 (1924); Sim,
Bryo. S. Afr. 161 (1926). Type species: H. perichaetiale (Hook.) Brid.
Plants medium to large, forming dense tufts; saxicolous or corticolous. Stems erect,
branching; central strand present. Leaves squarrose above erect bases. Costa short-excurrent;
in section round, with dorsal and ventral stereid bands. Upper laminal cells rounded-quadrate,
incrassate, mammillose ventrally; alar cells differentiated.
Perichaetial leaves very long, convolute; apices sometimes reaching capsule base. Capsule
cylindrical; peristome teeth 16, irregular, perforated or cleft to base.
Only 8 of the 60 species presently recognized in Holomitrium are found outside the tropics. The greatest
concentration of species is found at high altitudes in Central and South America. In Southern Africa Holo-
mitrium is restricted to the fynbos biome of the southern Cape and the alpine grass-heath biome of Lesotho and
western Natal, and the forests of northern Transvaal.
Holomitrium cylindraceum ( P . Beauv.)
Wijk & Marg. var. cucullatum ( Besch .)
Wijk & Marg. in Taxon 9: 190 (1960). Type:
Reunion, Menzies s.n.
Holomitrium vaginatum (Hook.) Brid. fo. cuculla-
tum Besch. in Annls Sci. nat. Bot., ser. 6, 9: 328
(1880). — var. cucullatum (Besch.) C. Mull., Gen.
Muse. Fr. 254 (1900).
Holomitrium affine Card. & Ther. in Bull. Soc.
bot. Geneve, ser. 2, 3: 249 (1911); Broth, in Natiirl.
PflFam. 10: 201 (1924); Sim, Bryo. S. Afr. 161
(1926). Syntypes: Natal, Medley Wood 712b
(NH1; H); Tanzania, Usambara, Holst 4215a, 2627a,
4209 (H; PC); Bourbon, Rodriques s.n. (PC); St
Philippe, Rodriques s.n. (PC); Mauritius, Robillard
s.n. (PC); Voeltz s.n. (PC).
Plants of medium size, in loose tufts,
dark green to light green above ; saxicolous or
corticolous. Stems erect, 10-20 mm tall,
frequently branched, tomentose below; in
section round, central strand present, small,
inner cortical cells of medium size, in 5-6
rows, outer cortical cells in single row, small,
incrassate, yellow-brown. Leaves circinate
above erect base dry, squarrose above base
Fig. 34. — Oreoweisia erosa (1-9): 1. habit, dry, x 1 ; 2. habit, wet, x 1 ; 3. habit, wet, x4; 4. stem in cross
section, x 130; 5. leaves, x 15; 6. leaf in cross section, x 300; 7. leaf base, x 100; 8. leaf apex, x 170; 9. part of
capsule mouth with rudimentary peristome teeth and spores, x 75. Holomitrium cylindraceum var. cucullatum
(10—19) : 10. habit, wet, x 1 ; 11. habit, dry, X 1 ; 12. habit, wet, x4; 13. habit, dry, x4; 14. stem in cross section
x 130; 15. leaves, x 10; 16. leaf in cross section, x 300; 17. cells at leaf base (right side), x 100; 18. leaf apex,
xlOO; 19. part of capsule mouth with peristome teeth and spores, X 120. (1-9, Sim 8047; 10—19, Crosby &
Crosby 8070).
130
Dicranaceae
wet, keeled; acuminate above an obovate
base, 2, 5^4,0 mm long; apex acute, cucul-
late, mucronate; margins erect, entire. Costa
short-excurrent; in section guide cells 2,
large, ventral substereid band 1 cell thick
(absent in distal section), ventral surface cells
slightly larger, incrassate, dorsal stereid band
2 cells thick, dorsal surface cells larger,
incrassate. Upper laminal cells quadrate to
rounded, or some short-rectangular, extend-
ing down margins of upper base, mammillose
ventrally; juxtacostal cells papillose dorsally;
basal cells oblong, becoming longer near
costa, longitudinal walls irregularly incras-
sate to weakly nodose, pitted; alar cells
enlarged, weakly thickened, yellowish.
Perichaetia terminal, leaves long con-
volute, oblong, abruptly short-acuminate,
8-10 mm long; leaf cells oblong throughout.
Seta 10-13 mm long, yellow; capsule erect,
urn cylindrical to ovate-cylindrical, 3, 0-3, 5
mm long, yellowish; peristome yellowish red,
teeth irregularly perforated or cleft to base,
finely ornate-papillose; operculum rostrate;
calyptra cucullate; spores round, 14-17 pm,
granulate. Fig. 34: 10-19.
Holomitrium cylindraceum var. cucullatum is
found in Africa south of the Sahara and the East
African Islands. In Southern Africa, the species
occurs on trees and rocks in forests of the southern
Map 46. — • Holomitrium cylindraceum var. cucullatum
and southwestern Cape, Drakensberg Escarpment of
western Natal and Soutpansberg of northern Trans-
vaal. Map 46.
Vouchers: Crosby & Crosby 8095; Owen 8;
Russell 2561; Thorne PRE-CH6433; Von Breitenbach
177; Wager PRE-CHI 5 14.
Fertile plants are easily identified by the long,
sheathing perichaetial leaves. The vegetative leaves,
broadly reflexed above the erect obovate base and
cucullate apex, will help to identify sterile plants.
The narrow, weakly nodose basal cells strongly con-
trast with the enlarged, yellowish alar cells. The
latter, however, are easily broken away when the
leaves are removed from the stem.
3. DICRANOLOMA
Dicranoloma (Ren.) Ren. in Rev. Bryol. 28: 85 (1901); Broth, in Natiirl. PflFam. 10: 207
(1924); Sim, Bryo. S. Afr. 162 (1926). Type species: not designated.
Plants large to robust, forming large, loose cushions; saxicolous. Stems erect, branching;
central strand present or absent. Leaves erect to spreading wet or dry, frequently secund.
Costa frequently weak; in section subround to oval, stereid bands present. Laminal cells
incrassate, smooth; alar cells distinct, enlarged, coloured; border cells narrow, hyaline.
Perichaetia lateral through innovation, leaves distinct. Seta erect; capsule curved or
erect; peristome cleft above, striate below; operculum rostrate; calyptra cucullate.
A genus of 114 species, Dicranoloma is best represented in Australasia. Of the 6 species recognized in
Africa, D. billardieri is the most widespread. In Southern Africa the genus is rare in forest of the fynbos and
montane forest biomes.
1 Leaf cells strongly incrassate, pitted; alar cells quadrate to short-rectangular 1. D. billardieri
1 Leaf cells : thin-walled, non-porous; alar cells oblong below, upper row quadrate 2.D. entabeniense
Fig. 35. — Dicranoloma entabeniense (1-8): 1. habit, X 1 ; 2. branch, dry, X 1 ; 3. habit, x3; 4. leaves, x 10;
5. leaf in cross section, x 170; 6. cells at leaf base (right side), x90; 7. lower laminal cells at margin, x220; 8.
upper laminal cells in subula, X 220. D. billardieri (9-17): 9. habit, xl; 10. habit, x 4; 11. stem in cross section,
1 10; 12. leaves, < 10; 13. leaf in cross section, x250; 14. cells at leaf base (right margin), x90; 15. upper
laminal cells, < 320; 16. perichaetial leaves, x5; 17. part of capsule mouth with peristome teeth, x55. (1-8,
Boitomley PRE-CH3381 ; 9-10, 12 & 14-17, Boucher 2209; 1 1 & 13, Crosby & Crosby 8140).
Dicranaceae
131
132
Dicranaceae
1. Dicranoloma billardieri ( Brid .) Par.,
Ind. Bryol. 2: 24 (1904); Broth, in Natiirl.
PflFam. 10: 209 (1924); Sim, Bryo. S. Afr.
162 (1926). Type: Australia, Billardier s.n.
Dicranum billardieri Brid. in Bot. Ztg, Regensburg
1: 214 (1802).
Plants large to robust, in loose cushions,
dark green to yellowish green, blackish below;
saxicolous. Stems erect, 60-70 mm high,
branched above; in section round, central
strand small, inner cortical cells large, thick-
walled, outer cortical cells in 1-2 rows,
stereids, reddish. Leaves erect-spreading to
secund; ovate-subulate, 4, 5-5,0 mm long;
base rounded; margin entire below, serrate
in subula, plane below, indexed above,
bordered from base to acumen by 1-2 rows
of narrow, elongated cells. Costa weak,
narrow, ending in apex or occasionally
absent in some leaves; in section subround,
guide cells 2, dorsal and ventral stereid
bands l(-2) cells thick, exposed. Upper
laminal cells elliptical, incrassate, pitted,
becoming longer toward base; basal cells
oblong, strongly incrassate, pitted; border
ceils long-linear, hyaline, smooth, in cross
section substereids; alar cells enlarged, red-
dish brown, quadrate to hexagonal or short-
rectangular, to 8 cells high; basal juxtacostal
cells reddish brown, oblong, incrassate,
pitted, not enlarged.
Map 47. — • Dicranoloma billardieri
X Dicranoloma entabeniense
Perichaetia terminal, leaves sheathing,
6-7 mm long, apex obtuse. Seta erect, 15-20
mm long, red; capsule arcuate, 3 mm long,
strumose, reddish; peristome teeth 16, orange,
vertically striolate or papillose, occasionally
perforated below, cleft to near middle, apical
segments striolate, yellowish; operculum
long-rostrate, 2 mm long; spores round,
12-15 pm. Fig. 35: 9-17.
Although widespread in the Southern Hemis-
phere, D. billardieri is restricted, in Southern Africa,
to the southern and southwestern Cape, primarily
around Table Mountain, and a few locations in the
eastern Transvaal. In Africa the species is found in
montane forest and ericaceous heath as far north as
Kilimanjaro in Tanzania (Bizot et al., 1978). Map 47.
Vouchers: Bews 8471; Boucher 2209; Crosby &
Crosby 8140; Goldblatt 1661 A; Howes 6; Vorster 765.
This species is separated from D. entabeniense by
strongly incrassate, pitted median leaf cells and
quadrate alar cells. In addition the habitats of the two
species are quite distinct.
2. Dicranoloma entabeniense Magill in
Mem. bot. Surv. S. Afr. 43: 3 (1979). Type:
Transvaal, Soutpansberg, Entabeni Forest,
Bottomley PRE-CH338 1 (PRE, holo. ! ; H ; L ;
MO; NY).
Plants large, in lax tufts; saxicolous.
Stems erect, to 60 mm tall, branched above;
in section round, central strand absent, inner
cortical cells large, becoming smaller, incras-
sate toward margin, red-brown, outer cortical
cells in single row, substereids, red-brown.
Leaves appressed with flexuose tips dry,
spreading wet; ovate to oblong, subulate
above, 5, 0-5, 5 mm long; base weakly
auriculate; margins entire, plane below,
convolute above; hyaline border extending
from base to near apex, 2-6 cells wide below.
Costa short-excurrent, toothed at tip; in
section oval, guide cells 6, ventral stereid
band 2 cells thick, exposed, dorsal cells
slightly smaller than guide cells, incrassate,
dorsal surface cells in single row, stereids.
Upper laminal cells smooth, subquadrate;
median cells shortly rectangular, extending to
base juxtacostally, arranged in ± longitudi-
nal rows; border cells long-linear, hyaline;
alar cells enlarged, reddish brown, long-
rectangular, extending to costa or cells near
costa narrow and not enlarged, upper row of
alar cells quadrate.
Sporophyte not known. Fig. 35: 1-8.
Dicranaceae
133
This species is presently known only from dry
forests of the northern Transvaal. Map 47.
Voucher: Type only.
The narrow costa, differentiated alar cells,
bordered leaves and smooth leaf cells of this species
are characteristic of Dicranoloma. It is distinct from
the other Southern African species, D. billardieri by
its non-porous, shortly rectangular leaf cells and
oblong alar cells.
Insufficiently Known Species
Dicranoloma nitidulum (C. Mull.) Par., Ind.
Bryol. edn 2, 2: 28 (1904). Basionym: Dicranum
nitidulum C. Mull, in Hedwigia 38: 88 (1899). Type:
Cape, Table Mountain, sin. coll., July 1883, Herb.
Schliephacke. The type has not been seen; however
Sim (1926) refers the species to D. billardieri (Brid.)
Par.
4. LEUCOLOMA
Leucoloma Brid., Bryol. Univ. 2: 218 (1827); Broth, in Natiirl. PflFam. 10: 209 (1924); Sim,
Bryo. S. Afr. 163 (1926). Type species: L. bifidum (Brid.) Brid.
Plants small to large, in loose tufts, green to glaucous-green ; saxicolous or corticolous.
Stems erect, to 60 mm high, sparsely radiculose; in section round, central strand absent.
Leaves flexuous or secund dry; lanceolate to ovate, setaceous or subulate; base weakly auri-
culate; margins frequently convolute, bordered throughout, wide below. Costa narrow, ending
in apex; in section oval, with dorsal and ventral stereid bands. Laminal cells short, strongly
papillose above; border cells long-linear, hyaline; intermediate cells smooth, rectangular;
juxtacostal cells variable; alar cells differentiated, enlarged and coloured.
Dioicous. Perichaetia terminal. Seta erect; capsule straight, cylindrical; peristome divided
to middle; operculum rostrate; calyptra cucullate.
The 131 species of Leucoloma are primarily tropical in distribution. The greatest concentration of species is
found on the East African Islands and in Central Africa. In Southern Africa Leucoloma is found on rock or
wood, in forests, from Table Mountain to Natal and the eastern and northern Transvaal.
The genus is divided from other members of the subfamily by its strong leaf border of hyaline cells and
papillose upper leaf cells. The leaves are also distinct in having as many as three other types of cells, e.g. differen-
tiated alar cells, intermediate cells lying between the border and upper leaf cells, and distinct basal leaf cells.
1 Papillose, quadrate laminal cells extending to base juxtacostal ly 1 . L. chrysobasilare
1 Basal cells rectangular, smooth, differing from laminal cells:
2 Median and lower leaf cells with several papillae scattered over lumen 2. L. sprengelianum
2 Median and lower leaf cells generally with a single stellate or spinose papilla over lumen, occasionally
a few cells with two papillae:
3 Leaves short, to 3,0 mm long, hyaline border narrow below, to 6 cells wide. . . .3. L. syrrhopodontioides
3 Leaves longer, 2, 5-6,0 mm long, hyaline border wide below, 12 or more cells wide 4. L. rehmannii
1. Leucoloma chrysobasilare (C. Mull.)
Jaeg., Adumbratio 2: 643 (1880); Broth, in
Natiirl. PflFam. 10: 210 (1924); Sim, Bryo.
S. Afr. 165 (1926). Syntypes: Comoro Islands,
Hildebrandt 1840 (BM!), 1842, 1846.
Dicranum chrysobasilare C. Mull, in Linnaea 40:
238 (1876).
Leucoloma zuluense Broth. & Bryhn in Forh.
VidenskSelsk. Krist. 1911 (4): 6 (1911); Broth, in
Natiirl. PflFam. 10: 210 (1924). Type: Natal, Zulu-
land, Ekombe, Titlestad s.n., July 1907, (H-BR!).
Plants large, in loose tufts, yellowish to
light green; corticolous or saxicolous. Stems
erect, 50-60 mm tall, occasionally branched;
in section central strand absent, inner cortical
cells in 5-8 rows, small, incrassate, outer
cortical cells in 2-3 rows of stereids or sub-
stereids, reddish. Leaves erect with tortuous
tips dry, erect spreading wet; ovate-subulate,
3, 5-5,0 mm long; apex weakly toothed;
margins plane, entire to serrate above,
bordered throughout by narrow, hyaline
cells, 2-6 cells wide below, 1-2 cells wide in
subula. Costa narrow, ending in apex or
rarely short-excurrent; in section guide cells
4, thickened, ventral and dorsal stereid band
2 cells thick, occasionally some cells sub-
stereids, surface cells undifferentiated. Lami-
134
Dicranaceae
nal cells quadrate to short-rectangular,
generally thickened, extending to leaf base
juxtacostally, bulging dorsally, with several
low papillae over lumen, ventral surface flat,
smooth; intermediate cells rectangular to
long-rectangular, smooth; alar cells distinct,
large, quadrate to short-rectangular, yellow-
ish brown, extending across entire leaf base,
quadrate near costa.
Immature perichaetial leaves similar to
vegetative leaves, papillose leaf cells not
reaching leaf base. Sporophyte not known.
Fig. 36: 11-17.
Originally described from the Comoro Islands,
L. chrysobasilare is now known from Madagascar
and the forests of East Africa and south through
Zimbabwe and Mozambique to the northern and
eastern Transvaal, Natal and Zululand. Map 48.
Map 48.— • Leucoloma sprengelianum
x Leucoloma chrysobasilare
Vouchers: Crosby <4 Crosby 7630, 7722; Magill
4914; Sim 236; Vorster 1537; Wager 54.
Leucoloma chrysobasilare is related to L. sprenge-
lianum but easily separated by the short, papillose
leaf cells extending to the leaf base on both sides of
the costa. In addition, in Southern Africa, L. chryso-
basilare is restricted to the northern and eastern
Transvaal. L. sprengelianum is most common in the
Cape, although a few collections have been made in
the eastern Transvaal.
The description of the sporophyte of L. chryso-
basilare by Sim (1926) was in error; the specimen
Wager 1 88 (as Sim 7728 PRE !) is L. rehmannii.
2. Leucoloma sprengelianum (C. Mull.)
Jaeg., Adumbratio 1: 115 (1871); Broth, in
Natiirl. PflFam. 10: 211 (1924); Sim, Bryo.
S. Afr. 164 (1926). Type: Prom, bonae spei,
Sprengel (BM !).
Dicranum sprengelianum C. Mull., Syn. Muse. 1 :
353 (1848).
Dicranum zeyheri C. Mull., Syn. Muse. 1 : 353
(1848). Leucoloma zeyheri (C. Mull.) Kindb., Enum.
Bryin. Exot. 64 (1888); Broth, in Natiirl. PflFam.
10: 211 (1924); Sim, Bryo. S. Afr. 163 (1926). Type:
Cape, Zeyher 496.
Plants medium-sized, in loose tufts,
yellowish green to green; terricolous or
saxicolous. Stems erect, 10-30 mm tall, little
branched; in section central strand absent,
inner cortical cells incrassate, in 8 rows,
smaller toward margin, outer cortical cells
in 1-2 rows, smaller, substereids, reddish.
Leaves erect to secund dry, spreading wet,
occasionally falcate; ovate to oblong, subu-
late, 3-4 mm long; apex acute, weakly
toothed; base weakly auriculate; margins
plane to erect, entire or serrulate above;
border of hyaline, elongated cells, 3-4 cells
wide at base, 1 cell wide above. Costa strong,
ending in subula; in section guide cells 4,
dorsal and ventral stereid bands 1-2 cells
thick, surface cells not differentiated. Laminal
cells quadrate or rounded-quadrate to short-
rectangular, dorsally papillose with numer-
ous low blunt papillae over lumen, occasion-
ally apical cells with only one or two stellate
or spinose papillae, laminal cells becoming
long-rectangular with seriate papillae toward
base; intermediate cells few, rectangular,
smooth, quickly merging with long-rectangu-
lar, incrassate basal cells; alar cells enlarged,
quadrate to rectangular, reddish brown.
Fig. 36. — Leucoloma sprengelianum (1-10): 1. habit, xl; 2. habit, xlO; 3-4. leaves, xl2; 5. leaf in cross
section, x 435; 6. cells at leaf base (right side), x 170; 7. lower laminal cells at margin, x 220; 8. leaf apex, xl70;
9. perichaetial leaf, X 1 2; 10. part of capsule mouth with peristome teeth, X 100. L. chrysobasilare (1 1—17):
11. habit, wet, x 1 ; 12. habit, dry, X 1 ; 13. upper stem, X 10; 14. leaves, X 12; 15. cells at leaf base (right side),
xl70; 16. upper laminal cells, x220; 17. leaf apex, Xl70. L. syrrhopodontioides (18-24): 18. habit, xl; 19.
habit, xlO; 20. leaves, x 12; 21. cells at leaf base (right side), x 170; 22. lower laminal cells at margin, x260;
23. upper laminal cells, X 260; 24. leaf apex, x 170. (1-10, Magill & Schelpe 4051 ; 1 1-17, Von Breitenbach 168;
18-24, Vorster 1452).
Dicranaceae
135
136
Dicranaceae
Perichaetial leaves sheathing, broadly
ovate, abruptly subulate, 3 mm long. Seta
straight, 9-10 mm long, reddish; capsule
short-cylindrical, to 1 mm long; peristome
reddish yellow, teeth divided above, occasion-
ally perforated below, vertically striate below,
papillose above; spores round, 25 jum,
weakly papillose. Fig. 36: 1-10.
Endemic to Southern Africa, L. sprengelianum
is collected on soil and rock in the fynbos biome of
the southern and southwestern Cape. A few collec-
tions have also been made in the forests of the eastern
Transvaal. Map 48.
Vouchers: Barnard PRE-CH3101; Cholnoky
1107; Schelpe 7825; Sim 9123; Wager PRE-CH381.
Some specimens have well-developed stellate
papillae on the leaf cells; but only a few cells will
have only single papillae. Even in these specimens
there is little difficulty with identification since the
lower leaf cells, near the junction with the inter-
mediate cells, are always rectangular with several
papillae per cell. Specimens of L. rehmannii and L.
syrrhopodontioides have quadrate or short-rectangular
lower leaf cells, with one or rarely two papillae per
cell.
Examination of the type of L. zeyheri shows that
it is conspecific with L. sprengelianum. Although older
collections labelled L. zeyheri are generally L.
sprengelianum, most recent collections, referred to
L. zeyheri, are small forms of L. rehmannii. The
species are similar in habit and, to some extent,
distribution; although L. rehmannii is more common,
especially in the Transvaal. The two species can be
easily separated on the basis of leaf cell ornamenta-
tion: L. sprengelianum ( —L. zeyheri) has numerous
low, blunt papillae scattered over the dorsal leaf
surface, and the papillae are seriate on cells just
above the base; in L. rehmannii each cell has a single,
large, stellate papilla on the dorsal surface. These
large papillae occur from just above the basal leaf
cells to the apex, where they become large, unbranched
spinose papillae. Although some variation occurs in
both species (vide descriptions) this character has
been effective in separating the two species.
3. Leucoloma syrrhopodontioides Broth.
in Bot. Jb. 24; 236 (1897); Broth, in Natiirl.
PflFam. 10: 211 (1924); Sim, Bryo. S. Afr.
163 (1926). Type: Transkei, Pondoland,
Bachmann 10 (H, holo. !).
Plants small, in loose tufts, yellowish
green; corticolous. Stems erect, to 15 mm
high, occasionally branched; in section central
strand absent, inner cortical cells in 4 rows,
incrassate, outer cortical cells in 1-2 rows of
substereids, reddish. Leaves flexuose dry,
erect-spreading wet; lanceolate-subulate, 2,5-
3,0 mm long; apex serrate; margin convolute,
entire below, border narrow, ending in
subula, to 6 cells wide below. Costa ending in
subula; in section guide cells 4, dorsal and
ventral stereid bands 2 cells thick, surface
cells not differentiated. Laminal cells
quadrate, dorsally papillose, each cell with
single, large, stellate papillae over lumen;
intermediate cells few, rectangular; basal
cells irregular, rectangular to rhomboidal,
incrassate; alar cells large, yellowish brown,
quadrate to short-rectangular, filling base.
Sporophyte not known. Fig. 36: 18-24.
Described from the Transkei, this species has
recently been collected in forests of the eastern Trans-
vaal. Map 49.
Map 49. — • Leucoloma rehmannii
x Leucoloma syrrhopodontioides
Voucher: Vorster 1452.
The plants are identified by their narrow border
in the leaf base, short leaves and unipapillose cells.
Small specimens of L. sprengelianum can be mistaken
for this species, but its pleuropapillose leaf cells help
to distinguish the two species.
4. Leucoloma rehmannii (C. Mull.) Rehm.
ex Par., Ind. Bryol. Suppl. 233 (1900); Broth,
in Natiirl. PflFam. 10: 211 (1924); Sim, Bryo.
S. Afr. 164 (1926). Type: Cape, Blanco,
Rehmann 29 (BOL!; PRE!).
Dicranum rehmannii C. Mull, in Hedwigia 38: 87
(1899).
Leucoloma haakonii Broth. & Bryhn in Forh.
VidenskSelsk. Krist. 1911(4): 6 (1911); Broth, in
Natiirl. PflFam. 10: 211 (1924). Type: Natal, Zulu-
land, Eshowe, Bryhn s.n., January 1909 (H-BR!).
Plants medium to large, in loose tufts,
light to yellowish green; corticolous or
saxicolous. Stems erect, 15-50 mm high; in
Dicranaceae
137
section central strand absent, inner cortical
cells in 3-4 rows, incrassate, outer cortical
cells in 1-2 rows, smaller, incrassate, reddish.
Leaves falcate to secund dry, erect-spreading
wet; lanceolate-subulate, 2,5-5,0(-6,0) mm
long; apex acute, toothed; margins weakly
convolute above base, entire below, serrate at
apex; border wide below, 1-2 cells wide to
apex. Costa percurrent; in section guide cells
4, dorsal and ventral stereid bands 2 cells
thick, surface cells undifferentiated. Laminal
cells quadrate above, rounded rectangular to
elliptical above base, dorsally papillose, with
l(-2) large, stellate papillae over lumen; inter-
mediate cells oblong; basal cells rounded-
rectangular to oblong-hexagonal; alar cells
enlarged, in 3-4 rows, rectangular below,
quadrate above, reddish yellow.
Perichaetial leaves with broad sheathing
base, abruptly long-subulate, 4,0-4, 5 mm
long. Seta red, 7, 0-7, 5 mm long; capsule
erect, urn cylindrical, 2, 0-2, 2 mm long;
peristome teeth cleft to middle, weakly
papillose, reddish yellow below, distal fila-
ments smooth, yellow; spores round, 17-20
nm, granulate. Fig. 37: 1-10.
Endemic to Southern Africa, L. rehmannii is
found on wood or rocks in forests of the southwestern,
southern and eastern Cape, Natal, Zululand, Swazi-
land and the eastern and northern Transvaal. Map 49.
Vouchers: Crosby & Crosby 8032, 9246; Junod
4013a; Schelpe 7851 ; Smook 866; Von Breitenbach 16;
Vorster 1430.
Leucoloma rehmannii is separated from other
Southern African species by its large, stellate papillae,
wide basal hyaline border and longer leaves.
The specimen ( H<f>eg 120, BM!) reported from
Southern Africa as L. albocinctum Ren. & Card.
(Dixon, 1932) belongs here. Examination of speci-
mens from the Cardot Herbarium (PC), indicates
Fig. 37. — Leucoloma rehmannii: 1. habit, wet,
x 1 ; 2. branch, dry, x 1 ; 3. habit, dry, x 1 ; 4. stem
in cross section, x 140; 5. leaves, xl2; 6. leaf in
cross section, x435; 7. cells at leaf base (right lower
margin), xl70; 8. laminal cells at right margin
showing border, x220;9. upper laminal cells, x350;
10. leaf apex, X170. (1—10, Von Breitenbach 159).
that L. rehmannii and L. albocinctum are similar;
however, further study is needed to assess the rela-
tionship between the two.
Many of the recent collections referred to L.
zeyheri belong here; see note under L. sprengelianum.
5. MICROCAMPYLOPUS
Microcampylopus (C. Mull.) Fleisch., Musci FI. Buitenzorg 1: 59 (1904); Broth, in Natiirl.
PflFam. 10: 183 (1924); Gangulee, Moss. E. India 269 (1971). Type species: M. subnanus
(C. Miill.) Fleisch.
Campylopus sect. Microcampylopus C. Mull, in Hedwigia 38: 77 (1899).
Plants small, caespitose; terricolous. Stems simple; central strand present. Leaves narrow,
linear-lanceolate to oblong-subulate; apex acute to obtuse; margins plane, entire to serrate
at apex. Costa narrow, excurrent as short, yellowish hair-point or filling subula; in section with
small ventral stereid or substereid band; dorsal stereids in small groups, dorsal surface rough.
Laminal cells quadrate to rectangular, incrassate; basal cells larger, rectangular; alar cells not
differentiated.
138
Dicranaceae
Dioicous. Perichaetia terminal. Seta flexuose or cygneous, yellowish; capsules ovate-
cylindrical; peristome teeth cleft to middle, vertically striate below, papillose above; operculum
rostrate; calyptra base fringed or entire; spores small.
A small genus of three species, Microcampylopus is found in Africa and the East African Islands, India,
Asia and Oceania. The single species recognized for Southern Africa, M. perpusillus, is found in the eastern
Transvaal, and extends northward into East Africa and the East African Islands. Microcampylopus nanus is
treated under Dicranella subsubulata.
Microcampylopus perpusillus {Mitt.)
Broth, in Natiirl. PflFam. 11: 525 (1925);
Dix. in S. Afr. J. Sci. 20: 314 (1923); Sim,
Bryo. S. Afr. 180 (1926). Type: Tanzania,
Ugogo, Hannington s.n. (BM, holo. !).
Campylopus perpusillus Mitt, in J. Linn. Soc., Bot.
22-: 301 (1886).
Campylopus angustinervis Dix. in S. Afr. ,T. Sci. 18:
303 (1922). Syntypes: Transvaal, Pietersburg, Junod
4001 (BM!); Belfast, Wager 884 (BM!; PRE!);
Zimbabwe, Matopos, Sim 8862, 8789, 8806 (BM!;
PRE!).
Plants small, loosely caespitose, green to
yellow-green above, brownish below; terri-
colous. Stems 3-6 mm high, simple; in section
round, central strand small, inner cortical
cells in 3-4 rows, outer row slightly smaller,
weakly thickened. Leaves appressed dry,
erect-spreading wet; linear-lanceolate, 2,5-
3,5 mm long; apex obtuse, weakly cucullate;
base scarcely differentiated, oblong; margins
broadly incurved or convolute dry, serrulate
at apex. Costa short-excurrent as yellowish,
smooth or denticulate awn, of width of
leaf base; in proximal section guide cells
6-10, ventral cells few, over central guide
cells, incrassate or substereids, dorsal stereids
in small groups separated by larger, incras-
sate cells, dorsal surface rough; in distal
section ventral cells absent or 1-4 over
central guide cells, dorsal stereid groups
separated by larger incrassate cells, dorsal
surface with single incrassate cells projecting
below stereid groups. Upper laminal cells
short-rectangular to rhomboidal, incrassate,
smooth; basal cells moderately differentiated,
slightly larger, rectangular, thin-walled; alar
cells not differentiated.
Sporophytes not known in Southern
Africa; described by Mitten as: Perichaetial
leaves undifferentiated. Seta flexuose; cap-
sule pendulous, buried, small, oval ; peristome
delicate, incomplete; operculum conic-ros-
trate; calyptra small, base fimbriate. Fig.
33: 1-8.
Rarely collected in eastern and Southern Africa,
M. perpusillus is found in woodlands and forests. In
the Flora area the species is only found in forests of
the northern and eastern Transvaal and southern
Natal. Map 44.
Vouchers: Magill 3400; Venter 4393.
The narrow, weakly convolute leaves, short-
excurrent costa and absence of differentiated alar
cells should separate M. perpusillus from related
species. It is somewhat similar to muticous forms of
Campylopus introflexus and differs from them mainly
in the size of plants, the development of the alar
cell region of the leaves and the stem anatomy.
6. CAMPYLOPUS
Campylopus Brid., Mant. Muse. 71 (1819); Broth, in Natiirl. PflFam. 10: 183 (1924); Sim,
Bryo. S. Afr. 165 (1926); Gangulee, Moss. E. India 275 (1971); Scott & Stone, Moss. S. Aust.
137 (1976); Smith, Moss FI. Brit. Irel. 166 (1978). Lectotype species: C. flexuosus (Hedw.)
Brid., vide Pfeiffer, Nom. 1 : 572 (1853).
Thysanomitrion Schwaegr., Spec. Muse. Suppl. 2: 61 (1823); Broth, in Natiirl. PflFam. 10: 188 (1924). Type
species: T. richardii (Brid.) Schwaegr.
Plants medium to robust, forming tufts, yellowish green to dark green; saxicolous or
terricolous. Stems erect, occasionally with white or reddish tomentum below; in section central
strand present. Leaves erect, rigid, narrow ; ovate to oblong or elliptical, subulate, channelled
to subtubulose above; margins entire to dentate in subula, not bordered. Costa broad, occu-
pying at least ^ of leaf base, frequently filling subula, percurrent to excurrent as long, frequently
dentate, hyaline awn; in section broad, guide cells numerous, medium-sized, thickened,
ventrally with either 1-2 layers of large, thin-walled cells completely covering guide cells or
Dicranaceae
139
with a stereid band 1-5 layers thick and generally not covering guide cells, dorsally with small
groups of stereids separated by larger incrassate cells or occasionally cells not differentiated,
dorsal surface smooth, rough or with lamellae 1-6 cells long below stereid groups. Laminal
cells quadrate, rectangular or rhomboidal, frequently incrassate; basal cells moderately
differentiated, somewhat larger, generally rectangular; alar cells generally differentiated, larger,
quadrate to rectangular, hyaline to reddish.
Dioicous. Perigonia gemmiform. Perichaetia terminal, leaves little differentiated. Seta
cygneous or arcuate ; capsule inclined, elliptical, sometimes curved, striate when dry and empty ;
peristome teeth 16, divided to middle, vertically striate below, orange-brown, papillose above,
hyaline; operculum rostrate; calyptra cucullate, fimbriate at base; spores finely papillose.
Approximately 600 species of Campylopus are currently recognized. The genus has a world-wide distribu-
tion with a concentration of species in tropical and subtropical areas. The major centres of described species
are South America and Africa. Forty-six species have been described or reported from the Flora area; only
twelve of these are recognized here.
1 Costa in proximal section with well defined ventral stereid or substereid band 1-3 cells thick, occasionally
ventral surface cells thin-walled:
2 Leaves 2-3 mm long; dorsal lamellae 2-4 cells long; costa percurrent or short-excurrent as smooth,
hyaline awn; frequently long-excurrent in comose leaves, and apical laminal cells hyaline
5. C. simii
2 Leaves (3-) 4-7 mm long; dorsal lamellae l(-2) cells long; costa percurrent or excurrent as dentate awn:
3 Leaves long-subulate; margins serrate above; costa percurrent or very short-excurrent, toothed:
4 In proximal section ventral surface cells differentiated, incrassate, occasionally interrupting 2-3
cell thick ventral stereid band; basal leaf cells weakly thickened, rarely pitted 4 . C. bequaertii
4 In proximal section costa without differentiated ventral surface cells, ventral stereid band 1(— 2)
cells thick; basal leaf cells strongly thickened, frequently irregularly so, strongly pitted
3. C. stenopelma
3 Leaves acuminate; margins entire or toothed at apex; costa short- or long-excurrent as a smooth to
dentate, hyaline or yellow awn :
5 Costa wider than \ width of leaf base; excurrent as a short, generally smooth awn
6. C. subchlorophyllosus
5 Costa less than f width of leaf base; excurrent as short, dentate awn:
6 Plants 40-80 mm high; quadrate basal leaf cells few; apical margins entire; costa excurrent as a
hyaline awn, rarely percurrent, in section with well defined stereid bands 1. C. ampliretis
6 Plants 10-30 mm high; quadrate basal leaf cells numerous; apical margins generally serrate;
costa excurrent as a yellow awn, in section with substereid bands 2. C. delagoae
1 Costa in proximal section with enlarged ventral cells in single layer, occasionally cells smaller, in 1-2
layers, incrassate:
7 Leaves long-setaceous, 7-12 mm long; dorsal costal surface smooth 12. C. procerus
7 Leaves acuminate to subulate, 2, 5-7,0 mm long; dorsal costal surface rough or with lamellae 1-6 cells
long:
8 Costa percurrent to short-excurrent, leaf tips occasionally hyaline; in proximal section dorsal costal
surface smooth to rough, rarely with projecting cells; in distal section dorsal surface smooth to
rough, rarely with lamellae 1-2 cells long:
9 Leaf margins serrate to serrulate above; basal leaf cells not extending further up margin than
costa :
10 Alar cells obvious, enlarged, reddish or infrequently hyaline 7. C. inchangae
10 Alar cells indistinct, not strongly differentiated from basal cells, occasionally hyaline and thin-
walled 8. C. pallidus
9 Leaf margins entire; basal cells generally hyaline, frequently extending further up margin than
costa, forming a weak V-shaped junction of basal and upper laminal cells:
1 1 In distal section, dorsal costal surface with lamellae 1-2 cells long 10. C. introflexus
11 In distal section, dorsal costal surface rough or with projecting cells 9. C. atroluteus
8 Costa excurrent as short or long, dentate awn ; in proximal section dorsal costal surface with pro-
jecting cells or single cell lamellae, in distal section dorsal surface lamellae 1-6 cells long:
12 Costa width of leaf base; dorsal costal lamellae 1-2 cells long 10. C. introflexus
12 Costa J width of leaf base; dorsal costal lamellae 2-6 cells long 11. C. pilifer
140
Dicranaceae
1. Campylopus ampliretis (C. Mull.) Par.,
Ind. Bryol. Suppl. 88 (1900); Broth., Naturl.
PflFam. 1 : 333 (1909). Type: Cape, Montagu
Pass, Rehmann 60 (PC), per. comm. Frahm;
vide Theriot in Dix. & Gepp (1923).
Dicranum ampliretis C. Mull, in Hedwigia 38: 81
(1899).
Dicranum olivaceonigricans C. Mull, in Hedwigia
38: 81 (1899). Campylopus olivaceonigricans (C. Mull.)
Par., Ind. Bryol. Suppl. 95 (1900). Type: Transvaal,
Duivels Krackler, Lydenburg, Wilms s.n., 1887
(G, holo. !).
Campylopus edwardsii Sim, Bryo. S. Afr. 168
(1926). Type: Transvaal, Johannesburg, Edwards
sub Sim 9836 (PRE, holo. !).
Thysanomitrion transvaaliense Herz. & Dix. ex
Sim, Bryo. S. Afr. 166 (1926). Type: Transvaal,
Graskop, Rolfes 188 (BM, holo.!, slide only).
Campylopus heteroneurum Ther. in Bull. Soc. bot.
Geneve 26: 79 (1936). Type: Transvaal, Sanatorium,
Junod s.n., 1900 (PC).
Plants large or robust, in loose turfs or
cushions, yellow-green to dark green above,
olive-green to blackish below; saxicolous or
rarely terricolous. Stems 40-80 mm tall, with
sparse, reddish tomentum below; in section
round, central strand large, inner cortical
cells of medium size, in 4-6 rows, outer
cortical cells in 2-3 rows, smaller, stereids to
substereids, reddish. Leaves frequently co-
mose at apex, and occasionally at intervals
along stem indicating previous growth cycles,
appressed dry, erect-spreading wet, convolute
or occasionally open, lamina frequently
weakly undulate; lanceolate to broadly
elliptical, narrowly acuminate, 3, 5-7,0 mm
long; base frequently weakly auriculate;
margins entire, broadly incurved above.
Costa frequently spurred in lamina, f width
of leaf base, short- to long-excurrent as a
dentate, hyaline awn or a very short hyaline
tip, rarely percurrent; in proximal section
guide cells c. 14, ventral stereid band 1-2
cells thick, central, exposing 1-2 guide cells
on either side, dorsal stereid band 2-3 cells
thick, occasionally cells substereids, dorsal
surface ± smooth; in distal section ventral
stereid band 1 cell thick, dorsal stereid band
2-3 cells thick, dorsal surface with single-
celled projections (lamellae). Upper laminal
cells rectangular to rhomboidal, rarely elon-
gate rhomboidal, generally strongly incrassate,
pitted; basal cells quadrate to rectangular,
in 2-6 rows above alar cells, thin-walled;
alar cells moderately to highly differentiated,
quadrate, weakly thickened, reddish to dark
red.
Sporophytes rare. Perichaetia terminal,
leaves slightly larger. Seta to 6 mm long,
blackish, frequently 2 per perichaetium ;
capsule elliptical, 2 mm long, plicate dry,
blackish; peristome fragile, only basal parts
seen; spores round, 20 pm, yellow-brown,
granulate. Other parts not seen. Fig. 38: 1-9.
Known from the Azores and Southern Africa,
C. ampliretis is frequently found on rock in forests or
woodlands of the Transvaal and Natal and occasion-
ally in forests of the southern Cape Province. Map 50.
Vouchers: Arnold 1271 ; Cholnoky 692, 800, 1079;
Hilliard & Bum 10467; Magill 3071, 5748, 5943;
Rankin 132; Von Breitenbach 57.
The size of the plants, broad leaves and presence
of a ventral stereid band in the costa will separate
this species from all other Southern African species
except C. delagoae (see note under that species).
A considerable degree of variation in the robust-
ness of individual specimens was observed. It was
found that this depended entirely on the extent to
which the leaves were convolute. Very robust plants
have open leaves, while less robust plants have con-
volute leaves. These differences could not be corre-
lated with any other characters of the plants, leaves or
areolation.
2. Campylopus delagoae (C. Mull.) Par.,
Ind. Bryol. Suppl. 91 (1900). Type: Trans-
vaal, near Otombi, between Delagoa Bay
and Lydenburg, Wilms s.n., 1884 (G, holo.!).
Dicranum delagoae C. Miill. in Hedwigia 38: 86
(1899). Thysanomitrion delagoae (C. Mull.) Broth, in
Naturl. PflFam. 10: 188 (1924).
Fig. 38. — Campylopus ampliretis (1-9): 1. habit, fertile and sterile stems dry, x 1 ; 2. habit, fertile and sterile
stems wet, x3; 3. leaves, x20; 4. leaf in proximal cross section (left half), x 170; 5. leaf in distal section (left
half), x 170; 6. cells at leaf base (right margin), x 170; 7. upper laminal cells, x 640; 8. upper juxtacostal cells,
X 170; 9. tip of awn, x 170. C. delagoae (10-17): 10. habit, xl; 11. habit, x3; 12. leaf, x 20; 13. leaf in proxi-
mal cross section (left half), x 170; 14. leaf in distal cross section (left half), x 170; 15. cells at leaf base (right
margin), Xl70; 16. upper laminal cells, x640; 17. tip of awn, xl70. C. stenopelma (18-25): 18. habit, xl;
19. habit, x3; 20 leaf, x20; 21. leaf in proximal cross section (left half), x 170; 22. leaf in distal cross section,
(left half), x 170; 23. cells at leaf base (right side), x 170; 24. upper laminal cells, x640;25. tip of subula, xl70.
(1-9, Smook & Phelan 854; 10-17, Magill 4562; 18-25, Smook & Phelan 875).
Dicranaceae
141
142
Dicranaceae
Dicranum inandae C. Mull, in Hedwigia 38: 95
(1899). Campylopus inandae (C. Miill.) Par., Ind.
Bryol. Suppl. 92 (1900). Thysanomitrion inandae
(C. Mull.) Broth, in Natiirl. PflFam. 10: 189 (1924).
Type: Natal, Inanda, Rehmann 43 (PRE!).
Plants medium to large, in loose cushions,
green to yellow-green above, blackish or
brownish below; terricolous or saxicolous.
Stems 10-30 mm tall, occasionally branched
below, with reddish tomentum below; in
section round, central strand small to me-
dium, inner cortical cells in 4-6 rows, yellow-
ish, outer cortical cells smaller, in 1-2 rows,
stereids to substereids, reddish. Leaves crowd-
ed, erect-appressed dry, patent wet; oblong
to lanceolate, acuminate, 3-5 mm long; base
squared; margins entire to serrate at apex,
erect to inflexed. Costa % width of leaf base,
short-excurrent, to 0,5 mm long, toothed,
yellowish; in proximal section guide cells
c. 12, ventral substereid band in single layer,
central, 2-3 guide cells exposed on both
sides, dorsal substereids in groups of 2-3
cells, substereid groups separated by single
incrassate cells, dorsal surface with single
projecting cell below each stereid group;
distal sections with similar anatomy, nar-
rower. Upper lamina l cells rhomboidal to
fusiform, weakly thickened, sinuolate; basal
cells quadrate to short-rectangular, weakly
thickened, quadrate at basal margins; alar
cells short-rectangular, thin-walled, reddish.
Sporophyte not known. Fig. 38: 10-17.
Campylopus delagoae is known from eastern and
Southern Africa. In the Flora area the species is found
Map 50. — • Campylopus ampliretis
X Campylopus delagoae
on rock or soil in forests or woodland of central and
eastern Transvaal, Swaziland and Natal. Map 50.
Vouchers: Cholnoky 503, 743; Hardy 4263;
Magill 3128, 3647; Smook 1460; Van Vuuren 1744;
Vorster 2152.
This species is very similar to C. ampliretis in
both morphology and Southern African distribution.
The two species are provisionally maintained in a
local flora context, on the basis of a few apparently
consistent character states. In C. delagoae the leaf
bases are squared and weakly bulging, and there is a
distinct region of quadrate basal leaf cells; laminal
cells are only weakly thickened and the costa in
section contains only substereid bands. On the other
hand, the leaf bases in C. ampliretis abruptly narrow
to the insertion and there are comparatively few
quadrate cells in the leaf base; laminal cells are
thickened, generally strongly so, and the costa, in
section, has well-developed stereid bands.
It is probable that these two species represent
forms of a single species, but a broader study, includ-
ing related species from the rest of Africa, is necessary
to properly assess the relationship.
3. Campylopus stenopelma (C. Mull.)
Par., Ind. Bryol. Suppl. 97 (1900); Broth,
in Natiirl. PflFam. 10: 188 (1924); Sim,
Bryo. S. Afr. 174 (1926). Syntypes: Cape,
Knysna, near Esternek, Rehmann 52 (NH!;
PRE!); Blanco, Rehmann s.n.
Dicranum stenopelma C. Miill. in Hedwigia 38:
83 (1899).
Dicranum chlorotrichus C. Mull, in Hedwigia 38:
87 (1899). Dicranodontium chlorotrichus (C. Mull.)
Par., Ind. Bryol. Suppl. 119 (1900). Campylopus
chlorotrichus (C. Miill.) Par., Ind. Bryol. edn 2, 1:
303 (1904); Broth, in Natiirl. PflFam. 10: 188 (1924);
Sim, Bryo. S. Afr. 172 (1926). Syntypes: Cape,
Montagu Pass, Rehmann 53 (BOL!; NH!; PRE!);
Knysna, Rehmann 53b (NH!; PRE!); Blanco,
Rehmann s.n.
Dicranum perfalcatum C. Miill. in Hedwigia 38: 87
(1899). Dicranodontium perfalcatum (C. Mull.) Par.,
Ind. Bryol. Suppl. 119 (1900). Campylopus capensis
Broth, in Natiirl. PflFam. 10: 188 (1924), non Cam-
pylopus perfalcatus Broth. (1901). Type: Cape, Knysna,
near Esternek, Rehmann s.n., 1875 (BM!; PC!).
Dicranum tenax C. Miill. in Hedwigia 38: 83 (1899).
Campylopus tenax (C. Mull.) Broth, in Natiirl.
PflFam. 10: 188 (1924). Type Cape, Blanco, Rehmann
54 (BOL!; NH!; PRE!).
Plants medium to large, in loose
cushions, yellow-green; terricolous or corti-
colous. Stems 10-25 mm tall, infrequently
branched, mainly below, lower stem covered
with reddish tomentum; in section subround
to elliptical, central strand small or large,
inner cortical cells in 4-6 rows, medium-
sized, incrassate, yellowish, outer cortical
cells substereids, in 2-4 rows, smaller, red-
dish. Leaves crowded frequently falcate.
Dicranaceae
143
erect to contorted dry, erect-spreading wet,
linear-lanceolate, subulate, 4-7 mm long;
base subauriculate ; margins erect to incurved,
entire below, serrate in upper subula. Costa |
of width of leaf base, percurrent; in proximal
section guide cells 14-20, ventral stereid
band small, in single layer, centred, exposing
3-4 guide cells on either side, dorsal stereids
in small groups, 2-3 cells each, stereid groups
separated by larger, incrassate cells; in distal
section ventral stereid band 1 layer thick,
centred, exposing 1-2 guide cells on either
side, dorsal cells incrassate or substereids,
dorsal surface with single cell lamellae.
Upper laminal cells rounded, quadrate to
short-rectangular or rhomboidal, thickened,
unpitted; basal cells rectangular, strongly
thickened, sometimes irregularly so, almost
smooth to strongly pitted ; alar cells enlarged,
quadrate, thin-walled, hyaline to reddish.
Sporophyte not seen. Fig. 38: 18-25.
Endemic to Southern Africa, C. stenopelma is
fairly widespread in woodland or forests. The species
is known from the central and eastern Transvaal,
Swaziland, Zululand, Natal, Lesotho, Transkei, and
the eastern, southern and southwestern Cape. Map 51 .
Map 51. — • Campylopus stenopelma
x Campylopus subchlorophyllosus
Vouchers: Cholnoky 679, 1090; Magill 3549,
6162; Oliver 7062; Smook 880.
The long, narrow, serrate leaves of this species
could be mistaken for those of similar species, such
as C. inchangae or C. stramineus. The presence of a
small ventral stereid band in the costa of the long,
narrow leaves will separate C. stenopelma from all
other Southern African species of Campylopus except
C. bequaertii. The latter two species are also easily
separated on the basis of costal anatomy.
4. Campylopus bequaertii Ther. & Nav.
in Bull. Soc. r. Bot. Belg. 60: 17 (1927);
Richards & Clear in Trans. Br. bryol. Soc.
5: 309 (1967). Type: Zaire, Ruwenzori,
Butagu, Bequaert 4974 (PC, holo.).
Plants medium to large, loosely caespi-
tose, green to yellow-green, blackish below;
terricolous. Stems 20-30 mm tall, branching
mainly above, with sparse, reddish tomentum;
in section round, central strand of medium
size, inner cortical cells in 5-6 rows, of
medium size, thin-walled, outer cortical
cells in 2-4 rows, thickened, red-brown.
Leaves widespreading, twisted dry, spreading
wet, linear-lanceolate, subulate, 4, 5-8,0
mm long; base weakly auriculate; margins
erect, serrate in subula. Costa f- of width of
leaf base, short-excurrent as toothed awn ; in
proximal section guide cells c. 18, ventral
stereid band strong, 2-3 cells thick, complete-
ly covering guide cells, ventral surface cells
in single layer across costa, incrassate,
occasionally interrupting stereid band, dorsal
stereid groups of 2-4 cells separated by
larger, thin-walled, triangular cells, dorsal
surface with a single, thin-walled cell pro-
jecting below each stereid group; in distal
section costa narrower, anatomy otherwise
similar. Upper laminal cells quadrate to
short-rectangular, weakly thickened ; basal
cells larger, rectangular to short-rectangular,
weakly thickened, not pitted; alar cells in
large distinct group, rectangular, thin-walled,
hyaline to reddish.
Map 52. — • Campylopus bequaertii
x Campylopus procerus
144
Dicranaceae
Sporophyte not known in Southern
Africa. Fig. 39: 1-8.
Campylopus bequaertii is known from western,
central and Southern Africa. In Southern Africa,
the species is known from Botswana, the northern,
central and eastern Transvaal, Swaziland, Natal and
the southwestern Cape. Map 52.
Vouchers: Cholnoky 746; Magill 3425, 4732;
Rankin 88.
The Southern African plants are more uniform
and perhaps distinct from the rather variable western
and central African element (Richards & Clear, 1967;
Bizot & Kilbertus, 1979). Specimens from the Flora
area are therefore only tentatively placed under this
species.
5. Campylopus simii Schelpe in Mem.
bot. Surv. S. Afr. 43: 5 (1979). Type: Orange
Free State, Kadziberg, Rehmann 58 (PRE,
holo.!).
Campylopus pseudojulaceus Sim, horn, illeg., Bryo.
S. Afr. 169 (1926), non C. Mull. (1898).
Plants small to medium, yellow-green;
terricolous. Stems 10-15 mm tall, occasional-
ly branched; in section round, central strand
large, inner cortical cells large, in 2-3 rows,
thin-walled, outer cortical cells small, in 1-4
rows, stereids to substereids. Leaves jula-
ceous, occasionally comose, appressed dry,
erect wet, oblong to elliptical, short-acumi-
nate, 2,0-2, 5 mm long; apex blunt, chloro-
phyllose; margins entire, incurved above,
plane below; comose leaves ovate to ligulate,
3 mm long; apex acuminate, hyaline. Costa $
of width of leaf base, percurrent, filling
acumen or occasionally very short-excurrent
as hyaline awn, in comose leaves long excur-
rent as hyaline awn; in proximal section
guide cells c. 24, ventral stereid band in
l(-2) row(s), sometimes substereids, centred,
exposing 3-5 guide cells on either side,
dorsal stereid cells in small groups of 3-4
cells each, stereid groups separated by larger,
thickened cells, dorsal surface with lamellae
below stereid groups, lamellae 2-4 cells long,
distal cells incrassate; in distal section costa
narrower, anatomy similar. Upper laminal
cells fusiform, thickened; basal cells highly
differentiated, rectangular to linear, thin-
walled, hyaline, extending further up margin
than costa, forming distinct V-shaped junc-
tion with laminal cells; alar cells weakly
developed, quadrate, thin-walled, hyaline to
reddish.
Sporophytes not known. Fig. 39: 9-16.
Endemic to Southern Africa, C. simii is presently
known only from rock at high altitude in the eastern
Orange Free State. Map 53.
Map 53. — • Campylopus inchangae
x Campylopus simii
Voucher : Type only.
The presence of a ventral stereid band in the
costa and long dorsal lamellae will separate C. simii
from the other Southern African species. The type
specimen is also distinct by the presence of comose
tufts of leaves with differentiated hyaline apices.
6. Campylopus subchlorophyllosus C.
Mull, ex Rabenh., Bryoth. Eur. 28: n. 1362
(1884). Type: Cape, Somerset East, Bosch-
berg, MacOwan s.n. (BM1; GRA!).
Fig. 39. — Campylopus bequaertii (1-8): 1. habit, x 1; 2. habit, x3; 3. leaf, x20; 4. leaf in proximal cross
section (left half), x 170; 5. leaf in distal cross section (left half), x 170; 6. cells at leaf base (left margin), x 170;
7. upper laminal cells, x640; 8. tip of subula, x 170. C. simii (9-16): 9. habit, xl; 10. habit, x3; 11. leaf,
x 20; 12. leaf in median cross section (left half), x 170; 13. cells at leaf base (right side), x 170; 14. upper laminal
cells, x640; 15. leaf apex, xl70; 16. leaf from comose head, x20. C. subchlorophyllosus (17-24): 17. habit,
x 1 ; 18. habit, x 3 ; 19. leaves, x 20: 20. leaf in proximal cross section, X 170; 21. leaf in distal cross section,
x 170; 22. cells at leaf base (right side), x 170; 23. upper laminal cells, x640; 24. leaf apex, x 170. C. pallidus
(25-32); 25. habit, x 1 ; 26. habit, x 5; 27. leaves, x 20; 28. leaf in proximal cross section, X 170; 29. leaf in distal
cross section, x 170; 30. cells at leaf base (right side), xl70; 31. upper laminal cells, x640; 32. leaf apex,
X 170. (1-8, Magill 4732; 9-16, Rehmann 58, 17-24, Cholnoky 1004; 25-32, Haggarth 4).
Dicranaceae
145
3C
146
Dicranaceae
Dicranum basalticolus C. Miill. in Hedwigia 38: 82
(1899). Campylopus basalticolus (C. Miill.) Par., Ind.
Bryol. Suppl. 89 (1900); Broth, in Natiirl. PflFam.
10: 187 (1924). Type: Cape, Somerset East, Bosch-
berg, MacOwan s.n., 1878 (BM!).
Plants small to medium, in loose
cushions, yellow-green, brownish below;
terricolous. Stems 5-15 mm tall, little
branched; in section round, central strand of
medium size, inner cortical cells in 4 rows,
thin-walled, outer cortical cells stereids or
substereids, in 1 row, reddish. Leaves crowd-
ed, appressed dry, erect-spreading wet;
oblong to elliptical, acuminate, 3-4 mm long;
base not differentiated; margins entire,
broadly incurved. Costa f-f of width of leaf
base, percurrent to excurrent as short,
smooth or dentate, hyaline awn; in proximal
section guide cells c. 24, small, thickened;
ventral stereid band 1-2 cells thick, complete-
ly covering guide cells, dorsal stereids in
small groups of 2-3 cells each, stereid groups
separated by large, incrassate cells, dorsal
surface with a single projecting cell below
each stereid group; in distal section ventral
stereid band 1-2 cells thick, central, exposing
3-4 guide cells on either side, dorsal stereid
groups smaller, of 1-2 cells each, separated
by large, incrassate cells, dorsal surface with
single-celled lamellae below each stereid
group, outer walls strongly thickened. Upper
laminal cells rounded, quadrate to short-
rectangular, incrassate; basal cells elongate,
irregularly shaped, elliptical to rectangular
or triangular, thickened, not pitted; alar
cells not strongly differentiated, quadrate,
thin-walled, reddish.
Sporophyte not known. Fig. 39: 17-24.
Endemic to Southern Africa, C. subchloro-
phyllosus is rarely collected on soil in fynbos or
grasslands of the southern and central Cape,
western Natal and eastern Transvaal. Map 51.
Vouchers: Cholnoky 725, 1004; Smook 1410.
Although C. subchlorophyllosus is not a very dis-
tinctive species, it does not appear to be closely
related to any other Southern African species. It is
identified by the presence of a small ventral stereid
band in the costa, short, narrow leaves, and dorsal
lamellae consisting of single projecting cells.
7. Campylopus inchangae (C. Miill.) Par.,
Ind. Bryol. Suppl. 92 (1900); Dix. in Trans.
R. Soc. S. Afr. 8: 185 (1920); Broth, in
Natiirl. PflFam. 10: 186 (1924). Type:
Natal, Inchanga, Rehmann 42 (BM!; PC!).
Dicranum inerangae C. Miill., in Hedwigia 38: 83
(1899) orth. var.; Campylopus imerangae (C. Miill.)
Par., Ind. Bryol. Suppl. 92 (1900), orth. var., vide
Dixon (1920).
Dicranum altovirescens C. Miill. in Hedwigia 38:
152 (1899). Campylopus altovirescens (C. Miill.) Par.,
Ind. Bryol. Suppl. 88 (1900); Broth, in Natiirl.
PflFam. 10: 186 (1924). Type: Natal, Jammerlappen,
Dittrich s.n., 1897 (H-BR!).
Dicranum longescens C. Mull, in Hedwigia 38: 85
(1899); Campylopus longescens (C. Mull.) Par., Ind.
Bryol. Suppl. 94 (1900); Broth, in Natiirl. PflFam.
10: 186 (1924). Type: Cape, Knysna, Esternek,
Rehmann 41 (NH!; PRE!).
Dicranum nanotenax C. Mull, in Hedwigia 38: 82
(1889). Campylopus nanotenax (C. Miill.) Par., Ind.
Bryol. Suppl. 94 (1900); Broth, in Natiirl. PflFam.
10: 186 (1924). Syntypes: Natal, Mapumulo, Borgen
s.n., 1867; Van Reenens Pass, Rehmann 51 (BM!;
G!).
Dicranum serridorsus C. Mull, in Hedwigia 38: 84
(1899). Campylopus serridorsus (C. Miill.) Par., Ind.
Bryol. Suppl. 97 (1900); Broth, in Natiirl. PflFam.
10: 185 (1924). Type: Cape, Table Mountain, Reh-
mann 45 (PRE!).
Dicranodontium laxitextum Broth. & Bryhn in
Forh. Vidensk Selsk. Krist. 1911 (4): 7 (1911).
Campylopus bryhnii Broth, in Natiirl. PflFam. 10:
188 (1924). Type: Natal, Zululand, Ekombe, Titlestad
s.n., 1907 (H-BR, holo.!; PC!).
Plants medium to large, in cushions,
green to yellow-green; terricolous or saxi-
colous. Stems 10-40 mm tall, frequently with
reddish tomentum below; in section round,
central strand small, inner cortical cells in
5-6 rows, of medium size, outer cortical
cells in 2-3 rows, smaller, reddish. Leaves
weakly contorted dry, spreading wet, linear-
lanceolate, subulate to setaceous, 4-7 mm
long; base auriculate; margins plane to erect,
serrate to serrulate above. Costa -J— \ of width
of leaf base, percurrent; in proximal section
guide cells c. 22, ventral cells large, thin-
walled, completely covering guide cells,
dorsal stereids in groups of 4-8 cells, stereid
groups separated by larger, strongly incrassate
cells, dorsal surface smooth to rough; in
distal section ventral cells incrassate, com-
pletely covering guide cells, dorsal stereid
groups of 4 cells separated by a larger
incrassate cell, dorsal surface with single-
celled lamellae under stereid groups. Upper
laminal cells small, quadrate to rhomboidal or
rectangular, incrassate; basal cells larger,
rectangular, incrassate, occasionally weakly
pitted; alar cells generally well defined,
quadrate, enlarged, reddish.
Dicranaceae
147
Perichaetia terminal, leaves oblong,
subulate to setaceous, 8-9 mm long. Seta 6-7
mm long, flexuose to cygneous, red-brown;
capsule arcuate, urn cylindrical, 1 ,5-2,0 mm
long; exothecial cells rectangular, incrassate;
peristome teeth 16, triangular, 0,5 mm long,
cleft above, striate below, papillose above,
reddish; operculum rostrate; spores round,
12-15 pm, irregularly papillose. Fig. 40:
14-21.
Campylopus inchangae is known from eastern
and Southern Africa. In the Flora area the species is
infrequently collected on soil or rock and occasionally
at the base of trees in grassland and woodland of
Transvaal, Swaziland, Zululand, Natal, Lesotho,
Transkei and the southern and southwestern Cape.
Map 53.
Vouchers: Cholnoky 243; Magill 3528, 4791,
4899, 5091, 6139; Rankin 284; Schelpe 7578.
Campylopus inchangae is most similar in general
appearance to C. stenopelma or C. bequaertii, but lacks
the stereid cells in the ventral costa. The ventral
costal cells of C. inchangae are large and thin-walled
in section and together with the small leaf cells and
long, narrow, serrate leaves, will help to place
specimens.
8. Campylopus pallidus Hook. f. & Wils.
in Hook, f., FI. Nov. Zel. 2: 68 (1854);
Scott & Stone, Moss. S. Aust. 142 (1976).
Syntypes: New Zealand, North Island, East
Coast and Auckland, Colenso s.n., Sinclair
s.n.
Dicranum pulvinatus C. Miill. in Hedwigia 38: 80
(1899), horn, illeg. Campylopus pulvinatus ( C . Mull.)
Par., Ind. Bryol. Suppl. 96 (1900); Sim, Bryo. S. Afr.
179 (1926), horn, illeg. Type: Cape, Stinkwater,
Rehmann 62 (PRE !).
Plants of medium size, forming cushions,
yellow-green to light green; terricolous or
saxicolous. Stems 2-15 mm high, branching
mainly above, with reddish tomentum below;
in section round, central strand small, cells
collapsed, inner cortical cells in 3-4 rows,
thin-walled, outer cortical cells in 2-3 rows,
incrassate, reddish. Leaves crowded, tips
weakly contorted dry, erect-spreading wet;
linear-lanceolate to ovate-subulate, 2, 5-6,0
mm long, broadly channelled ventrally;
margins broadly incurved, entire to serrulate.
Costa | of width of leaf base, percurrent
or excurrent as short, denticulate tip; in
proximal section guide cells c. 18, ventral
cells large, completely covering guide cells,
dorsal stereids in groups of 4 to 15 cells,
stereid groups separated by larger incrassate
cells or occasionally cells undifferentiated and
uniformely incrassate, dorsal surface cells
often larger, incrassate, dorsal surface rough;
in distal section ventral cells incrassate,
completely covering guide cells, dorsal cells
incrassate or stereids, dorsal surface with pro-
jecting cells or single-celled lamellae. Upper
laminal cells quadrate to rhomboidal, incras-
sate; basal cells rectangular, thin-walled,
hyaline to yellowish; alar cells not differen-
tiated to weakly bulging, rectangular, thin-
walled, hyaline to yellowish or rarely reddish.
Perichaetia terminal, leaves undifferen-
tiated. Seta 10-12 mm long, yellowish;
capsule cylindrical, 0,8-1 ,2 mm long, plicate
dry; exothecial cells rectangular, sinuate,
incrassate; peristome teeth 16, triangular,
cleft above, 0, 3-0,4 mm long, striate below,
papillose and hyaline above; operculum
rostrate; calyptra cucullate, 1,5 mm long;
spores round, 13-15 pm, pale yellow, granu-
late. Fig. 39: 25-32.
Widespread in the Southern Hemisphere, C.
pallidus has only recently been recognized from South-
ern Africa. In the Flora area the species is most
commonly collected in Natal; however, specimens are
frequently collected in the montane grasslands or
forests of Transvaal, eastern Orange Free State,
Lesotho, Zululand and the southern and south-
western Cape. Map 54.
Map 54. — • Campylopus pallidus
Vouchers: Magill 5071, 5732; Oliver 7355; Van
Rooy 457 ; Von Breitenbach 270.
The species is related to C. inchangae ; the two
species differ primarily in the development of alar
cells and the dentation of the subula of the leaves.
Campylopus pallidus is also similar to some facies of
C. atroluteus from which it is most easily separated
148
Dicranaceae
by: (1) quadrate laminal cells that extend nearly to
the apex, (2) weakly serrulate margins of the subula,
and (3) basal cells not or only rarely extending higher
up the margin than the costa, a condition that is
common in C. atroluteus in Southern Africa.
9. Campylopus atroluteus (C. Mull.) Par.,
Ind. Bryol. Suppl. 89 (1900); Broth, in
Natiirl. PflFam. 10: 187 (1924); Sim, Bryo.
S. Afr. 178 (1926). Syntypes; Cape, Stink-
water, Rehmann 44 (NH!); Cape Town,
Rehmann 63 (BOL!; NH!; PRE!).
Dicranum atroluteus C. Mull, in Hedwigia 38: 80
(1899).
Dicranum hartramiaceus C. Mull, in Hedwigia 38:
86 (1899). Campylopus hartramiaceus (C. Miill.)
Ther. ex Broth, in Natiirl. PflFam. 10: 185 (1924).
Type: Cape, Cape Town, Rehmann 37 (PRE!).
Dicranum catarractilis C. Mull, in Hedwigia 38:
79 (1899). Campylopus catarractilis (C. Miill.) Par.,
Ind. Bryol. Suppl. 90 (1900); Broth, in Natiirl.
PflFam. 10: 187 (1924); Sim, Bryo. S. Afr. 179
(1926). Type: Cape, Devil’s Peak, Rehmann 64 (PRE!).
Dicranum leptotrichaceus C. Miill. in Hedwigia
38: 84 (1899). Campylopus leptotrichaceus (C. Miill.)
Par., Ind. Bryol. Suppl. 93 (1900); Broth, in Natiirl.
PflFam. 10: 185 (1924); Sim, Bryo. S. Afr. 172(1926).
Type: Cape, Knysna, Esternek, Rehmann s.n., 1875
(BM!).
Dicranum ampliretis C. Miill. in Hedwigia 38: 81
(1899). Type: Cape, Montagu Pass, Rehmann 60
(quoad specimen herb. BM!).
Plants of medium size, loosely caespi-
tose, yellow-green, brownish below; terri-
colous. Stems 10-20 mm high, with sparse,
reddish tomentum below; in section round,
central strand large, inner cortical cells in
3-4 rows, thin-walled, outer cortical cells in
2 rows, smaller, thickened, reddish. Leaves
weakly contorted dry, erect-spreading wet;
ovate-lanceolate to elliptical, subulate, 4-6
mm long; apex occasionally with hyaline
tip; base scarcely differentiated; margins
entire, inrolled above. Costa f of width of
leaf base, percurrent or short-excurrent,
yellow to hyaline; in proximal section guide
cells c. 20, ventral cells large, thin-walled or
occasionally incrassate, dorsal cells undif-
ferentiated, incrassate or strongly differen-
tiated medianally, with small groups of
stereids separated by larger, incrassate cells,
laterally without stereid groups, all cells
large, forming multistratose area 1-8 cells
wide, dorsal surface smooth to rough; in
distal section costa frequently occupying
entire leaf, ventral cells smaller, incrassate,
completely covering guide cells, dorsal cells
frequently undifferentiated, stereids or sub-
stereids, dorsal surface rough or with single-
celled projections. Upper laminal cells
quadrate to rectangular or elongate, sinuo-
late, incrassate; basal cells frequently reach-
ing higher along margins than costa, narrowly
rectangular, thin-walled; alar cells scarcely
differentiated, occasionally reddish.
Sporophyte not known in Southern
Africa. Fig. 40: 1-13.
Found throughout Africa and the West African
Islands, C. atroluteus is generally collected on soil in
grassland and woodland. In Southern Africa the
species is frequently collected in the southern and
southwestern Cape, Natal, Lesotho, Zululand, eastern
Orange Free State, Swaziland, and Transvaal. A few
specimens have also been collected in the eastern
Cape and the Caprivi Strip of South West Africa/
Namibia. Map 55.
Vouchers: Cholnoky 278, 380; Emdon 3; Magill
5791, 5803, 6344; Smook 829; Van Rooy 573.
An unusually complex species, C. atroluteus is
made up of several entities that could not be satis-
factorily separated, except at the extremes of each
group. Therefore a rather broad species concept is
accepted here (see also Frahm, 1976) encompassing a
large amount of variation between individual speci-
mens. The variation is most frequently expressed in
leaf size and shape, apex shape and anatomy of the
costa.
Fig. 40. — Campylopus atroluteus (1-13): 1. habit, x 1 ; 2. habit, xl; 3. habit, x 1 ; 4. habit, x2; 5. leaves,
x 20; 6. leaf in proximal cross section (left half, typical form), x 170; 7. leaf in proximal cross section (left half,
intermediate form), X 170; 8. leaf in proximal cross section (right half, ‘ hartramiaceus’ facies), x 170; 9. leaf
in distal cross section (right half, typical form). X 170; 10. cells at leaf base (right side), x 170; 11. junction of
basal and laminal cells, xl70; 12. upper laminal cells, X640; 13. leaf apex, Xl70. C. inchangae (14-21): 14.
habit, x 1 ; 15. habit, x2; 16. leaf, X20; 17. leaf in proximal cross section (left half), X 170; 18. leaf in distal
section (left half), X 170; 19. cells at leaf base (left margin), X 170; 20. tip of subula, X 170; 21. laminal cells
in upper subula, x435. C. procerus (22-33): 22. habit, xl; 23. leaf, x20; 24. leaf in proximal cross section
deft half), x 170; 25. leaf in distal cross section (left half), X 170; 26. cells at base of leaf (left margin), x 170;
27. upper laminal cells, x 170; 28. tip of subula, X 170; 29. calyptra, x20; 30. capsule, x5; 31. deoperculate
capsule and calyptra, X 5; 32. part of capsule mouth with peristome tooth, x90; 33. enlargement of peristome
tooth, x 640, (1, Vorster 448a; 2 & 7, Lavranos 15297 a; 3-6 & 9-13, Cholnoky 919; 8, Rehmann 37; 14-21,
Rehmann 42; 22-33, Wager 209).
Dicranaceae
149
150
Dicranaceae
Map 55. — • Campylopus atroluteus
One of the most interesting variations is expressed
in the costal anatomy of facies bartramiaceus. Small
dorsal stereid groups are present in the central part
of the costa, but laterally the cells are undifferen-
tiated. The cells are similar in size and shape to the
ventral cells, thus giving the leaves a multistratose
appearance up to 8 cells wide on either side. A similar
development, although involving only 1-3 rows of
cells, has been seen in other specimens of C. atrolu-
teus and occasionally in C. inchangae and C.
introflexus.
The South African specimens referred to C.
bicolor (C. Mull.) Wils. (Sim, 1926; Dixon, 1920;
Frahm, pers. comm.) differ in leaf shape and costal
anatomy from the Australian and New Zealand
specimens that have been examined. These specimens
{Hall 4, 5, 6, BM!; Van Zanten 7608198, PRE!) are
therefore provisionally treated as a facies of the
species complex treated here as C. atroluteus. The
Southern African specimens, that have been referred
to C. bicolor, have leaves that are narrowly subulate
above an oval or ovate base. The leaf apices are
frequently narrow and cucullate, although occasional-
ly some leaves have straight apices or even short,
hyaline tips. The leaves of the isosyntypes of C.
atroluteus have a similar shape and most apices are
straight with short, hyaline tips. A few of the leaves,
however, have narrow, cucullate apices. Another
specimen {Garside 6283) with a slightly longer,
denticulate, hyaline point, has recently been referred
to C. bicolor var. ericeticola (C. Mull.) Dix. (pers.
comm. Frahm, 1980).
Dixon (1920) went into the subject of variation in
leaf apices of this group in some detail, when he
recorded C. bicolor from Southern Africa. He also
suggested a possible relationship to C. pseudobicolor.
The relationship of this Malagasy species and South-
ern African species must still be explored.
In view of the plasticity of this ‘species’ as well
as its facies that occasionally resemble forms of C.
introflexus or C. pallidus, and its apparent relation-
ship to C. bicolor, the above specimens are provision-
ally treated under the species complex C. atroluteus.
The problem of further separation of this complex
into recognizable taxa, whether at the specific or
subspecific level, will only be solved when the problem
is examined on a much broader scale.
10. Campylopus introflexus ( Hedw .)
Brid., Mant. Muse. 72 (1819); Broth, in
Natiirl. PflFam. 10: 187 (1924); Sim, Bryo.
S. Afr. 170 (1926); Gangulee, Moss. E.
India 300 (1971); Scott & Stone, Moss.
S. Aust. 140 (1976); Smith, Moss FI. Brit.
Irel. 183 (1978). Type: Nova Hollandia.
Dicranum introfiexum Hedw., Spec. Muse. 147
(1801).
Dicranum lepidophyllus C. Miill., Syn. Muse. 1 :
407 (1848). Campylopus lepidophyllus (C. Miill.)
Jaeg., Adumbratio 1 : 139 (1872); Sim, Bryo. S. Afr.
171 (1926). Syntypes: Cape, Palmiet River, Ecklon
s.n. (BM!); Caledon, Gnetpagar, Ecklon s.n.
Dicranum leucobasis C. Miill. in Hedwigia 38: 78
(1899). Campylopus leucobasis (C. Miill.) Par., Ind.
Bryol. Suppl. 93 (1900); Dix. in Trans. R. Soc. S.
Afr. 8: 184 (1920). Type: Cape, Montagu Pass,
Rehmann 71 (NH!; PRE!).
Dicranum weisiopsis C. Miill. in Hedwigia 38: 79
(1899). Campylopus weisiopsis (C. Miill.) Par., Ind.
Bryol. Suppl. 99 (1900); Broth, in Natiirl. PflFam. 10:
184 (1924). Type: Cape, Table Mountain, Rehmann
61 PC!).
Campylopus echinatus Sim, Bryo. S. Afr. 175
(1926), nom. illeg. Syntypes: Cape, Rehmann 67, 68,
69, 70, 71, 72; Sim 8581b, 8604; Natal, Sim 9837
(PRE!).
Plants medium to large, in cushions,
yellow-green to green above, frequently with
reddish tomentum below; terricolous. Stems
(5-)10-20 mm tall, rarely branched; in
section round, central strand large, inner
cortical cells in 3-5 rows, large, thin-walled,
outer cortical cells in 1-3 rows, smaller,
substereids, red-brown. Leaves frequently
comose at stem apex and occasionally at
intervals along older stems, appressed dry,
erect-spreading wet, oblong to elliptical,
acuminate, 3-7 mm long; base scarcely
differentiated; margins incurved above, entire.
Costa of width of leaf base, long excur-
rent up to 2 mm as a hyaline, dentate awn,
frequently reflexed, or infrequently percur-
rent, apex with short hyaline tip; in proximal
section guide cells 27-30, ventral cells in
single layer, large, weakly thickened, dorsal
stereid cells in small groups of 2-4 cells,
occasionally substereids, stereid groups separ-
ated by large, incrassate cells, dorsal surface
with single large, incrassate cells below
stereid groups; in distal section ventral cells
similar, dorsal cells generally substereids
Dicranaceae
151
throughout, dorsal surface with lamellae,
1-2 cells long. Upper laminal cells short-
rectangular to rhomboidal or quadrate;
basal cells moderately differentiated, reaching
higher along margin than costa, rectangular
to elongate, thin-walled, hyaline; alar cells
variably differentiated, short-rectangular,
thin-walled, reddish.
Sporophyte not known from Southern
Africa (Gradstein & Sipman, 1978). Fig.
41:1-11.
Campylopus introflexus is presently recognized
from southern South America, Southern Africa,
Australia and New Zealand, and has been introduced
into England and Europe. In Southern Africa the
species is known from fynbos, grasslands and wood-
lands of the Cape, Natal, Zululand, Lesotho, Orange
Fig. 41. — Campylopus introflexus (1-11): 1. habit, X 1; 2. habit, x 1; 3. habit, x3; 4. leaf, x20; 5. leaf in
proximal cross section (left half), xl70; 6. leaf in distal cross section (left half), xl70; 7. cells at leaf base
(right side), x 170; 8. junction of basal and laminal cells, x 170; 9. tip of awn, x 170; 10. basal leaf cells, xl70;
11. upper laminal cells, X 640. C. pilifer (12-20): 12. habit, x 1 ; 13. habit, xl; 14. habit, x3; 15. leaf, x20;
16. leaf in proximal cross section (left half), x 170; 17. leaf in distal cross section (right half), x 170; 18. cells at
leaf base (right side), x 170; 19. upper laminal cells, x640; 20. tip of awn, x 170. (1-11, Magill 6060; 12-20,
Cholnoky 962).
152
Dicranaceae
Free State, Transvaal and from a few collections from
southeastern Botswana and central South West
Africa/Namibia. Although uncommon throughout its
range in Southern Africa, the species is more fre-
quently collected in the southern and southwestern
Cape. Map 56.
Vouchers: Cholnoky 419, 727 ; Jacot Guillarmod
8104; Hardy 4264; Lavranos 15207c; Magill 4415,
6060; Schelpe 7916; Van Rooy 71.
The extreme variability of C. introflexus in most
of its distinguishing characters makes identification
of some specimens very difficult. In its typical form,
with comose tufts of leaves at the stem apex and long,
hyaline, reflexed awns, the species is easily recog-
nized, but only about 40% of the Southern African
specimens have this combination of characters. Over
50% of the specimens examined did not have reflexed
awns or in some cases lacked awns altogether.
A considerable degree of variation in the develop-
ment of stereids in the costa was observed throughout
the range of the species. The smaller plants fre-
quently show weak development of stereid groups.
Many of the forms of C. introflexus in Southern
Africa approach the habit of C. pilifer. Several recent
publications (vide Frahm, 1974; Gradstein & Sip-
man, 1978) have discussed the differences between the
two species. Southern Africa is one of the few areas
where the species are sympatric. In addition to the
characters used in the key, C. pilifer is more common
in the northern parts of the Flora area, while C.
introflexus is more common in the south. See also
notes under C. pilifer.
11. Campylopus pilifer Brid., Mant.
Muse. 72 (1819); Frahm in Rev. bryol.
lich. 40: 33-44 (1974); Gradstein & Sipman
in Bryologist 81: 114-121 (1978). Type:
Italy, ‘In Insula Ischia ad rupes, Auguste
1806’ (B, lecto.).
Campylopus polvtrichoides De Not., Syll. Muse.
222 (1838); Broth, in Naturl. PflFam. 10: 187 (1924).
Type: Europe.
Campylopus purpurascens Lor., Moosstud. 158
(1864); Broth, in Naturl. PflFam. 10: 187 (1924); Sim,
Bryo. S. Afr. 176 (1926). Type: Cape, Table Moun-
tain, Ecklon s.n. (NY, holo. !).
Campylopus trichodes Lor., Moosstud. 159 (1864);
Broth, in Naturl. PflFam. 10: 187 (1924); Sim, Bryo.
S. Afr. 168 (1926). Type: Cape, Table Mountain,
Ecklon s.n. (NY!).
Dicranum griseolus C. Miill. in Hedwigia 38: 80
(1899). Campylopus griseolus (C. Miill.) Par., Ind.
Bryol. Suppl. 92 (1900); Broth, in Naturl. PflFam.
10: 187 (1924); Sim, Bryo. S. Afr. 167 (1926).
Type: Transvaal, Lydenburg, Wilms s.n., 1887 (G,
holo.!).
Dicranum purpureoaureus C. Miill. in Hedwigia 38 :
82 (1899). Campylopus purpureoaureus (C. Miill.)
Par., Ind. Bryol. Suppl. 96 (1900); Broth, in Naturl.
PflFam. 10: 187 (1924). Syntypes: Transvaal, Duivels
Krackler, near Lydenburg, Wilms s.n., 1887 (G);
Liebenbergsvley, Rehmann 59 (PRE!).
Campylopus trichodes var. perlamellosus Dix. in S.
Afr. J. Sci. 18: 304 (1922); Sim, Bryo. S. Afr. 168
(1926). Syntypes: Zimbabwe, Rhodes’s Grave,
Matopos, Sim 8858 (BM!; PRE!); Natal, Tugela
River, Bews sub Sim 8374; (BM! ; PRE!).
Campylopus bewsii Sim, Bryo S. Afr. 176 (1926).
Syntypes: Natal, Zwartkop, Sim 9838 (PRE!);
Mont-Aux-Sources, Bews sub Sim 9891 (PRE!);
Giant’s Castle, Symons s.n. (PRE!).
ICampylopus symonsii Sim, Bryo. S. Afr. 167
(1926). Syntypes: Natal, Giant’s Castle, Symons sub
Sim 9843, 9835 (PRE!).
Plants large to robust, forming large
turfs or cushions, green to yellow-green
above, black-green below; terricolous or
saxicolous. Stems (5-) 10-40 mm tall, infre-
quently branched; in section round, cen-
tral strand small, inner cortical cells in 6
rows, of medium size, weakly thickened,
outer cortical cells in 2-4 rows, stereids,
reddish. Leaves crowded, rarely comose at
stem apex, elliptical to ovate-oblong, acumi-
nate to subulate, (3-)4-7 mm long; base
auriculate to scarcely differentiated; margins
entire, erect to incurved above. Costa of
width of leaf base, short-excurrent as denticu-
late, hyaline awn 0, 2-2,0 mm long; in
proximal section guide cells c. 30, ventral
cells in single layer, large, thin-walled, rarely
thickened, centred, exposing 2-4 guide cells
on either side, dorsal stereids in small groups,
3-4 cells each, stereid groups separated by
larger incrassate cells, occasionally all dorsal
Dicranaceae
153
cells similar, strongly incrassate, dorsal
surface with lamellae under stereid groups,
2-4 cells long; in distal section costa nar-
rower, ventral cells somewhat smaller, occa-
sionally strongly incrassate, dorsal anatomy
similar, dorsal lamellae 4-6 cells long. Upper
laminal cells quadrate to angular, incrassate,
median laminal cells rectangular, rhomboidal
or elongate, shapes generally mixed, incras-
sate; basal cells moderately differentiated,
reaching higher along margin than costa,
rectangular, thin-walled, hyaline; alar cells
quadrate, thin-walled, reddish or hyaline.
Sporophytes not known in Southern
Africa (Gradstein & Sipman, 1978). Fig.
41: 12-20.
Widespread in Europe, Asia, the Americas and
Africa, C. pilifer is most common in grasslands and
woodlands of the Transvaal, Swaziland, Zululand,
Natal, Lesotho and Orange Free State. Specimens
have also been occasionally collected in the eastern
and western Cape. Map 57.
Vouchers: Cholnoky 285, 501; Esterhuysen
20224; Hilliard & Burn 10408; Jacobsz 5010; Magill
3364, 5593; Pienaar 20; Smook 1425.
Although similar to C. introflexus, the present
species is generally separated by the longer dorsal
costal lamellae. In addition C. pilifer only rarely has
comose tufts (heads) of leaves at the stem apex and
never has reflexed awns.
Specimens of C. symonsii are here considered only
a facies of C. pilifer. The specimens, which are fairly
widespread in upper elevations of the Drakensberg,
could, however be separated by their very long, some-
what recurved, serrate hair-points, and spurred costa
with an undivided dorsal stereid or substereid band.
12. Campylopus procerus (C. Mull.) Par.,
Ind. Bryol. 258 (1894); Broth, in Natiirl.
PflFam. 10: 186 (1924). Type: Tanzania,
Mt Kilimandjaro, Meyer s.n., 1889 (BM!;
PC!).
Dicranum procerus C. Mull, in Flora, Jena 73:
472 (1890).
Plants large to robust, loosely caespitose,
yellow-green above, blackish below; saxi-
colous. Stems 30-90 mm tall, branching
below, frequently with reddish tomentum
below; in section round, central strand very
small, inner cortical cells in 5-6 rows,
medium-sized, yellowish, outer cortical cells
in 2-3 rows, smaller, stereids, dark red.
Leaves twisted, curved dry, widely spreading
to recurved wet, oblong to elliptical, seta-
ceous, 7-12 mm long; base scarcely differen-
tiated, rounded; margins inrolled to erect
above, denticulate to dentate above. Costa f
of width of leaf base, percurrent; in proximal
section guide cells 46-60, small, ventral cells
large, lax, completely covering guide cells,
dorsal stereid groups small, of 2-3 cells,
stereid groups separated by larger incrassate
cells, dorsal surface flat; in distal section
ventral cells smaller, dorsal stereid groups
distinct, dorsal surface flat. Upper laminal
cells small, quadrate, rectangular or rhom-
boidal, incrassate; basal cells weakly dif-
ferentiated, large, rectangular to rhomboidal,
incrassate, pitted; alar cells differentiated,
thin-walled, reddish.
Perichaetia terminal, polysetaceous,
leaves convolute, oblong-setaceous, 8-12 mm
long. Seta flexuose to cygneous, to 10 mm
long, yellowish to red-brown ; capsule arcuate,
plicate dry; urn elliptical, to 2 mm long,
strumose; exothecial cells rectangular, incras-
sate; peristome teeth 16, triangular, 0,4 mm
long, cleft above, striate below, papillose,
hyaline above; operculum rostrate, 1,3 mm
long; calyptra cucullate, 2,5 mm long,
fringed; spores round, 12—17 pm, weakly
papillose. Fig. 40: 22-33.
Known from central, western and Southern
Africa, C. procerus is infrequent throughout its range.
In Southern Africa the species is rarely collected on
rock in forests of the eastern Transvaal. Map 52.
154
Dicranaceae
Vouchers: Crosby & Crosby 13401; Smook &
Phelan 845 ; Vorster 486a.
Stems and leaves of C. procerus are considerably
longer than those of any of the other Southern African
species. In addition the very prominent ventral cells
of the costa give the leaf sections a distinctive ap-
pearance.
Insufficiently Known Species
Campylopus aureoviridis (C. Miill.) Schimp. ex Par.,
Ind. Bryol. Suppl. 89 (1900). Basionym: Dicranum
aureoviridis C. Miill. in Hedwigia 38: 88 (1899).
Type: Cape, Houteniqua, Breutel s.n., 1862. The type
has not been seen; however Sim (1926) refers the
species to C. atroluteus (C. Miill.) Par.
7. CHORISODONTIUM
Chorisodontium (Mitt.) Broth, in Nattirl. PflFam. 10: 204 (1924); Bell in Br. Antarct. Surv.
Bull. 37: 33 (1973); Robinson in Smiths. Cont. Bot. 27: 24 (1975). Type species: not desig-
nated.
Dicranum sect. Chorisodontium Mitt, in J. Linn. Soc., Bot. 12: 62 (1869).
Plants large, caespitose; saxicolous. Stems erect; central strand present. Leaves erect to
falcate-secund, lanceolate-subulate, frequently subconvolute; margins serrate to entire, without
borders. Costa broad, occupying £ width of leaf base, filling subula; in section guide cells
numerous, dorsal and ventral stereid bands strong, dorsal surface cells vertically elongate,
occasionally projecting dorsally as large sharp mammillae. Laminal cells rectangular to linear,
incrassate, occasionally pitted; basal cells rectangular, thin-walled; alar cells greatly enlarged,
reddish.
Dioicous. Perichaetia terminal, leaves little differentiated. Seta long, straight; capsules
erect, cylindrical, slightly asymmetrical; peristome teeth cleft to middle, striolate below;
operculum long-rostrate; calyptra cucullate, base entire; spores often large.
The 14 species of Chorisodontium are found in the Subantarctic Islands, Africa and South America north-
ward along the Andes to Central America. The following new species is the first report of the genus for the
African continent. The genus is similar to Campylopus ; the two genera are most easily separated on costal
anatomy, leaf cell shape and ornamentation as well as several sporophyte characters.
Chorisodontium falcatum Magill, sp. nov.,
habitu C. aciphyllo (Hook.f. & Wils.) Broth,
et C. dicranellato (Dus.) Roiv. simile, sed
foliis falcato-secundis, subula valde serrata,
pagina dorsale costae mammillosissima et
anatomia costae differt.
Type: Cape, Hex River Mountains,
Roodeberg, on rock, Esterhuysen 20972
(BOL, holo. ; H; L; MO; NY; PRE).
Plants large, caespitose, green to yellow-
green above, brownish below; saxicolous.
Stems 30-70 mm tall, occasionally branching,
with sparse, reddish or whitish tomentum
below; in section round, central strand small,
inner cortical cells large, in 4-6 rows, outer
cortical cells small, weakly thickened, in 2-3
rows. Leaves somewhat crowded, falcate-
secund with flexuose subulae dry, falcate-
secund but less flexuose wet, subulate to
setaceous above oblong to oval base, 6-9 mm
long; ventral surface broadly channelled
above; margin erect, strongly serrate or
occasionally almost entire. Costa broad, ^
of width of leaf base, filling subula, ventral
superficial cells linear, smooth, dorsal super-
ficial cells rectangular, becoming shorter
above base, sharply mammillose; in section
ventrally concave, guide cells (20-) 30-34(-50)
above base, medium-sized, strongly thick-
ened; in proximal section ventral stereid
band strong, 2-4 cells thick, ventral surface
cells not differentiated, dorsal stereid band
strong, 3-4 cells thick, dorsal surface cells
not differentiated; in distal section ventral
substereid band 2 cells thick centrally,
quadrate, laterally cells in single layer,
quadrate, incrassate, ventral surface cells not
differentiated, dorsal internal cells in 2
layers, slightly smaller than guide cells,
Dicranaceae
155
absent laterally, dorsal surface cells vertically
elongate, incrassate, projecting dorsally as
large, sharp mammillae, tip of mammillae
strongly thickened. Laminal cells rectangular,
incrassate, frequently irregularly thickened,
pitted, smooth; in section strongly thickened,
lumens horizontally compressed; basal cells
becoming long-rectangular, thin-walled,
linear at basal margin; alar cells large,
broadly rectangular, thin-walled, reddish.
Sporophyte not known. Fig. 42: 1-12.
The species is presently known from rock recesses
and shaded ledges at the base of cliffs at upper eleva-
tions in the mountains of the southwestern Cape.
The species has also been collected on Kerguelen
Island {Engel 7334, 7345, 7355 (ALTA), comm. D. H.
Vitt). Map 58.
Map 58. — • Leucobryum acutifolium
x Chorisodontium falcatum
Vouchers: Esterhuysen 16827, 21334, 27048,
27864; Wilms 2567 (Gl).
This species is similar to C. aciphyllum (Hook,
f. & Wils.) Broth, and C. dicranellatum (Dus.) Roiv.
but differs in the falcate, secund leaves, strongly
serrate subulae, coarsely mammillose dorsal costa
and costal anatomy.
Fig. 42. — Chorisodontium falcatum: 1. habit, wet,
x 1 ; 2. habit, dry, x 1 ; 3. habit, x 3 ; 4. stem in cross
section, xlOO; 5. leaf, x20; 6. leaf in proximal
cross section (left half), x 170; 7. leaf in median cross
section (left half), x 170; 8. leaf in distal cross section
(left half), xl70; 9. cells at leaf base (right margin),
xl70; 10. juxtacostal cells (dorsal surface), x640;
11. upper laminal cells, x640; 12. tip of subula,
X 170. (1—1 2, Esterhuysen 20972).
156
Dicranaceae
Subfamily Leucobryoideae
Plants small to large, in dense cushions, glaucous-green; terricolous or saxicolous. Stesm
erect, with or without central strand. Leaves thickened, consisting mostly of broad, multi-
layered costa; in section with single row of small, rhombic chlorocysts surrounded by 2-8
layers of large, hyaline leucocysts. Laminal cells large, hyaline, confined mostly to leaf base.
Perichaetia terminal. Seta erect; capsule erect, frequently cylindrical, strumose; peristome
teeth cleft to middle, vertically striate below, papillose above; operculum rostrate; calyptra
cucullate.
Only a single genus of this subfamily, Leucobryum, is represented in Southern Africa.
The family Leucobryaceae is not recognized here. The genera frequently treated in Leucobryaceae are
gametophytically similar; however, sporophyte characters, especially peristome architecture, clearly illustrate
the heterogeneity of the family. The genera of Leucobryaceae have therefore been divided between the
families Dicranaceae and Calymperaceae (fide Crosby & Magill, 1977). Only two Southern African genera
are involved: Leucobryum with a dicranaceous peristome is placed in Dicranaceae and Octoblepharum is moved
to Calymperaceae.
LEUCOBRYUM
Leucobryum Hampe in Linnaea 13; 42 (1839); Broth, in Natiirl. PflFam. 10: 223 (1924); Sim,
Bryo. S. Afr. 181 (1926). Type species: L. glaucum (Hedw.) Schimp.
Plants medium to large, forming large cushions, whitish to glaucous-green; terricolous
or saxicolous. Stems branching below, central strand present or absent. Leaves crowded,
thickened, fleshy, lanceolate to oval-ligulate; apex acute to obtuse; lamina narrow, restricted
to base, hyaline. Costa filling upper leaf; in section leucocysts in 2-8 layers, enclosing ±
median chlorocysts. Laminal cells quadrate, rectangular or fusiform, thin-walled, hyaline.
Dioicous. Dwarf male plants growing on or among leaves of female plants. Perichaetia
terminal. Seta erect, elongate; capsule inclined, asymmetric, 8-ribbed dry, strumose; peristome
teeth 16, triangular, deeply cleft, vertically striate below; operculum long-rostrate; calyptra
cucullate.
Leucobryum contains 124 species and is found in tropical and temperate regions throughout the world.
In Southern Africa the genus is infrequently collected in forests or dense woodlands.
1 Plants large; leaves in distal section with leucocysts in 5-8 rows 3. L. madagassum
1 Plants medium; leaves in distal section with leucocysts in 2 rows;
2 Leaves 3-6 mm long, spreading wet 1. L. acutifolium
2 Leaves 7-10 mm long, generally falcate wet 2. L. rehmannii
1. Leucobryum acutifolium {Mitt.) Card.
in Bull. Herb. Boissier, ser. 2, 4: 105 (1904);
Broth, in Natiirl. PflFam. 10: 224 (1924).
Syntypes: Tanzania, Usagara Mountains,
Hannington s.n.; Last s.n.; Natal, Saunders
s.n.; Madagascar, Metier s.n. (all NY!).
Schistomitrium acutifolium Mitt, in J. Linn. Soc.,
Bot. 22: 302 (1886); Sim, Bryo. S. Afr. 183 (1926).
Leucobryum gueinzii C. Miill. in Hedwigia 38: 58
(1899); Broth, in Natiirl. PflFam. 10: 224 (1924);
Sim, Bryo. S. Afr. 183 (1926). Syntypes: Cape,
Montagu Pass, Rehmann 74 (PRE!); Natal, Gueinzius
s.n.
Fig. 43. — Leucobryum madagassum (1-8): 1. habit, x 1 ; 2. habit, x 5; 3. leaf, x 20; 4. leaf in proximal cross
section, x70; 5. leaf in distal cross section, x70; 6. cells at leaf base (left side), x 170; 7. cells at leaf margin,
x 640; 8. leaf apex, Xl70. L. acutifolium (9-16) : 9. habit, X 1 ; 10. habit, X 5; 11. stem in cross section, Xl70;
12. leaf, x20; 13. leaf in proximal cross section, x70; 14. leaf in distal cross section, x70; 15. cells at leaf
margin, x 640; 16. leaf apex, X 170. L. rehmannii (17-24): 17. habit, X 1 ; 18. habit, X 5; 19. leaf, X 20; 20. leaf in
proximal cross section, x 70; 21. leaf in distal cross section, x 70; 22. cells at leaf margin, x 170; 23. upper leaf
cells, X640; 24. leaf apex, X 170. (1-8, Wager PRE-CH12038; 9-16, Rehmann 74; 17-24, Rehmann 75).
Dicranaceae
157
158
Dicranaceae
Plants small, loosely caespitose, glau-
cous-green; corticolous or saxicolous. Stems
5-10 mm tall, unbranched; in section round,
central strand very small, weak, inner cortical
cells in 4-5 rows, thickened, outer cortical
cells in 1-2 rows, smaller, reddish. Leaves
contorted dry, spreading wet, elliptical to
oblong, 3-6 mm long; lamina narrow, uni-
stratose; apex acuminate, apiculate; margins
entire, broadly incurved above. Costa broad;
in proximal section chlorocysts small, rhom-
boidal, ventral leucocysts in l(-2) layer(s),
dorsal leucocysts in 1 layer medianly, 2-3
layers laterally; in median and distal sections
ventral and dorsal leucocysts in single layers.
Upper laminal cells in 9-12 rows on either side
of costa, rectangular to quadrate, infre-
quently some cells triangular, thin-walled,
pitted; marginal cells rectangular to linear,
narrower above, extending to apex, hyaline.
Perichaetia terminal, leaves convolute,
acuminate, 4 mm long. Seta 10-15 mm long,
reddish; capsule curved, striated dry, stru-
mose, urn 1,0-1, 5 mm long; peristome red-
yellow, teeth 16, triangular, cleft to below
middle, 0,5 mm long; vertically striate
below, papillose above; operculum long-
rostrate, 1,0-1, 5 mm long; calyptra cucul-
late, 2,5 mm long, smooth; spores round,
15-17 pm, papillose, green. Fig. 43: 9-16.
This species is known from east Africa, the East
African Islands and Southern Africa. In the Flora
area, L. acutifolium is infrequently collected in moun-
tain or escarpment forests of the northern and
eastern Transvaal, Swaziland, and lowland forests of
Zululand, Natal, and the southern and southwestern
Cape. Map 58.
Vouchers: Cholnoky 663; Crosby & Crosby
9206; Kemp 807; Kluge 1054; Magill 3815; Oliver
7061 ; Rankin 144; Schelpe 7861.
Leucobryum acutifolium is separated from the
other Southern African species by its smaller size,
shorter leaves and leucocysts in distal leaf section in
two rows.
Theriot (1929) reported L. isleanum Besch. for
Southern Africa, from a Junod specimen, collected
in the northern Transvaal. A duplicate of the speci-
ment (G!) was examined and appears to be L.
acutifolium. The author could not locate the type of
L. isleanum among the specimens from the Bescherelle
collection (BM), so its relationship to L, acutifolium
could not be assessed. Leucobryum isleanum is
provisionally excluded from the Flora.
Leucobryum parvulum Card, was reported (Bizot,
1967) from Mt Anderson near Sabie in the eastern
Transvaal. One of the syntypes ( Perrot 86, PC!) was
examined; it is very similar to L. acutifolium. A more
comprehensive study must be undertaken to confirm
the relationship between these two species. The
species is provisionally excluded from the Flora.
2. Leucobryum rehmannii C. Mull, in
Hedwigia 38: 58 (1899). Type: Cape, Knysna,
Esternek, Rehmann 75 (BOL!; NH!; PRE!).
Leucobryum perfalcatum Sim, Bryo. S. Afr. 182
(1936), horn, illeg. Type: Knysna, Esternek, Rehmann
75 (PRE!; BOL!; NH!).
Leucobryum gracilifolium Dix. in K. norske Vidensk.
Selsk. Skr. 1932(4): 4 (1932). Syntypes: Cape,
Knysna, Deep Walls Forest Station, H4>eg 95, 102
(BM!).
Plants small to medium, in cushions,
glaucous-green, tan below; terricolous or
corticolous. Stems 10-20 mm tall, irregularly
branched below; in section round, central
strand present, inner cortical cells in 4-5
rows, incrassate, outer cortical cells in 2
rows, smaller, red-brown. Leaves crowded, ±
contorted dry, spreading to frequently falcate
wet; elliptical, abruptly acuminate, 7-10
mm long; lamina narrow, unistratose; base
scarcely differentiated; margins entire, plane
below, inrolled above. Costa broad; in
section chlorocysts small, rhomboidal, at
base ventral leucocysts in 1 row medianly,
l(-2) row(s) laterally, dorsal leucocysts in 1
row medianly, 2-3 rows laterally, in upper
proximal and distal sections ventral and
dorsal leucocysts in single layers. Upper
laminal cells in 6-7 rows on either side of
costa, irregularly rectangular to fusiform,
very thin-walled, pitted, hyaline.
Sporophyte not known. Fig. 43: 17-24.
Endemic to Southern Africa, L. rehmannii has
been collected rarely in the evergreen forests of the
southern Cape Province. Map 59.
Vouchers: Crosby & Crosby 7986, 9245; Russell
2522.
This species is very closely related to L. acuti-
folium and may represent only a robust form of that
species. Leucobryum rehmannii is provisionally main-
tained here because of the consistently larger plants,
longer, generally falcate leaves, and longer laminal
cells.
The distribution area of L. rehmannii is very
small when compared with the African distribution
area of L. acutifolium. Both species occur in the
southern Cape, but as yet are not known from the
same forests.
3. Leucobryum madagassum Besch. in
Annls Sci. nat. Bot., ser. 6, 9: 337 (1880);
Broth, in Natiirl. PflFam. 10: 224 (1924);
Sim, Bryo. S. Afr. 182 (1926). Type: Mada-
gascar, Rosas s.n., 1876 (BM ! ; PC!).
Dicranaceae
159
Plants large to robust, in cushions,
glaucous-green; saxicolous. Stems 10-40 mm
tall, irregularly branched below; in section
round, central strand absent, inner cortical
cells medium-sized, yellowish, outer cortical
cells in 1-2 rows, slightly smaller, red-brown.
Leaves appressed dry, erect-spreading wet,
broadly lanceolate to oblong, 5,0-7, 5 mm
long, concave; lamina narrow, unistratose;
apex acuminate; base scarcely differentiated,
rounded; margins entire, broadly incurved
above. Costa broad; in proximal section
chlorocysts small, rhomboidal, ventral leuco-
cysts in 2 layers, infrequently small areas with
3 layers, dorsal leucocysts in 2(-3) layers; in
distal section chlorocysts rhomboidal, ven-
tral leucocysts in 2(-3) layers, narrowing to 1
layer marginally, dorsal leucocysts in 3(-4)
layers, abruptly narrowing at margin. Upper
laminal cells in 6-9 rows on either side of
costa, rectangular, thin-walled, pitted; mar-
ginal cells in 2-3 rows, linear, hyaline, extend-
ing to apex.
Sporophyte not known in Southern
Africa. Fig. 43: 1-8.
Leucobryum madagassum is known from east
Africa, the East African Islands and Southern Africa.
Map 59. — • Leucobryum madagassum
x Leucobryum rehmannii
In Southern Africa the species has been collected in
forests of the northern and eastern Transvaal and
Swaziland. Map 59.
Vouchers: Cholnoky 586; Magill 3505; Stirton
6980; Vorster 2144.
Leucobryum madagassum is considerably larger
than the other two Southern African species. In
addition, the leaf in distal section has leucocysts in
5-8 layers; in both of the other species the leucocysts
are in 2 rows.
161
CALYMPERACEAE
Plants small to medium, in loose tufts or cushions; terricolous, saxicolous or corticolous.
Stems erect; central strand present or absent. Leaves elongate above broad, sheathing, ±
hyaline base; margins frequently thickened, toothed, or with hyaline border of elongated
cells, occasionally with intermarginal bands (teniolae) of elongated cells. Costa strong, per-
current or excurrent, occasionally gemmiferous apically. Upper laminal cells small, papillose;
frequently inner basal cells (cancellinae) enlarged, hyaline; sometimes leaves modified, having
chlorocysts surrounded by several layers of leucocysts. Gemmae multicellular, cylindrical,
frequently produced at leaf apices.
Seta erect, elongate; capsules, erect, urn ± cylindrical; peristome teeth 8, 16 or absent;
calyptra large, cucullate or campanulate, sometimes persistent.
A small family of 14 genera with tropical to temperate distributions.
1 Leaves several cells thick above; chlorocysts small, triangular, surrounded by large
leucocysts 1 . Octoblepharum
1 Leaf lamina unistratose above, margins frequently thickened :
2 Stem with small central strand; laminal cells not extending down margin
4. Hypodontium
2 Stem without central strand ; laminal cells extending down margin or between margin
and cancellinae :
3 Leaves with cartilaginous border of narrow, elongated, hyaline cells. . . .3. Syrrhopodon
3 Leaves without border of elongated cells, teniolae occasionally present:
4 Leaf margins serrate, denticulate or smooth 2. Calymperes
4 Leaf margins strongly and irregularly dentate, spinose at shoulders. . . 3. Syrrhopodon
1. OCTOBLEPHARUM
Octoblepharum Hedw., Spec. Muse. 50 (1801); Broth, in Nattirl. PflFam. 10: 225 (1924); Sim,
Bryo. S. Afr. 183 (1926). Type species: O. albidum Hedw.
Plants glaucous, loosely caespitose. Stems short, erect. Leaves strap-shaped above oval
base; multistratose above, small triangular chlorocysts surrounded by large leucocysts.
Perichaetial leaves undifferentiated. Seta short, erect; capsule exserted, short; peristome
short, teeth 8; calyptra cucullate.
A genus of 16 species of tropical and subtropical distribution, Octoblepharum is rare in the northern and
eastern parts of the Flora area.
The leaf anatomy indicates that the upper, multistratose part of the leaf is a modified costa and the oval,
basa 1 region is the lamina.
The family Leucobryaceae (see note under Dicranaceae subfamily Leucobryoideae) is not recognized in the
Flora. The genus Octoblepharum should, however, be segregated at the subfamily level (Octoblepharoideae)
from the rest of Calymperaceae in the Flora area. The family Leucophanaceae, recognized ‘primarily by posses-
sing the leucobryoid leaf’ (vide Edwards, 1980) is also not taken up here, because of the rather heterogeneous
leaf morphology exhibited by the group. A great deal of research is needed to properly assess the relationship of
the genera with ‘leucobryoid leaves’.
162
Calymperaceae
Calymperaceae
163
Octoblepharum albidum Hedw., Spec.
Muse. 50 (1801); Broth, in Natiirl. PflFam.
10: 226 (1924); Sim, Bryo. S. Afr. 183 (1926).
Type: Bahamas.
Plants small to medium, loosely caespi-
tose, glaucous-green; corticolous. Stems 2-4
mm high, unbranched; in section round,
central strand absent, cortical cells large, in
5 rows, outer row slightly thickened. Leaves
squarrose above base wet or dry, oblong
above a wider, oval base, 4-5 mm long;
apex rounded, abruptly apiculate; margins
plane, entire or serrulate at apex. Costa
filling upper leaf; in distal section reniform,
chlorocysts small, triangular, ventral leuco-
cysts in 3 layers, dorsal leucocysts in 2 layers.
Laminal cells quadrate; basal cells rectangu-
lar; leucocysts thin-walled, with large lateral
pores. Gemmae or rhizoidal growth frequent
at leaf apices, gemmae clavate, 4-6 cells
long.
Autoicous. Perigonia scattered along
stem; perichaetia terminal, leaves undif-
ferentiated. Seta 3-4 mm long, straight;
capsule small, erect, urn ovoid, ± 1 mm long,
symmetrical; peristome short, teeth 8, trian-
gular, 0, 1-0,2 mm long, yellow-brown;
operculum rostrate, beak oblique; calyptra
cucullate, deciduous; spores round, 17-20
pm, granulose. Fig. 44: 1-9.
Octoblepharum albidum is widespread in the
Americas, Africa and Australasia. In Southern Africa
the species is rarely collected at the base of trees or on
thick humus in closed forests of the central and
eastern Transvaal, Swaziland, Zululand and Natal.
Map 60.
Map 60. — • Octoblepharum albidum
x Calymperes rabenhorstii
Vouchers: Kemp 808; Schelpe PRE-CH12638;
Wager PRE-CH444.
The species is easily identified by its thick,
leathery, glaucous-green leaves and small size. The
upper leaves are bent sharply back above the appres-
sed base giving the plants a pressed or flattened
appearance.
2. CALYMPERES
Calymperes Sw. in Web., Tab. Exh. Calyptr. Operc. Gen. (1813); Broth, in Natiirl. PflFam.
10: 236 (1924); Sim, Bryo. S. Afr. 261 (1926); Edwards in J. Bryol. 11: 55 (1980). Lectotype
species: C. lonchophophyllum Schwaegr., vide Williams in Bull. Torrey bot. Club 47: 385.
(1920).
Plants dull, dark green, scattered or in tufts; corticolous. Stems short; central strand
absent. Leaves linear above hyaline sheathing base; margins thickened; teniolae often present.
Costa strong, percurrent to short-excurrent, frequently gemmiferous. Laminal cells small,
cancellinae nearly filling base.
Perichaetia terminal. Seta short; capsule cylindrical; peristome absent; calyptra campanu-
late, plicate, ± twisted, persistent.
Fig. 44. — Octoblepharum albidum (1-9): 1. habit, xl; 2. habit, x5; 3. leaves, xl2; 4. leaf in proximal
cross section, x75; 5. leaf in distal cross section, x75; 6. leaf cells at shoulder, xl30; 7. upper leaf cells at
margin, x 130; 8. leaf apex, x 130; 9. part of capsule mouth with peristome teeth and spores, x 1 10. Calymperes
levyanum (10—18) : 10. habit, wet, x 1 ; 11. habit, dry, xl; 12. habit, x4; 13. leaves, x 12; 14. leaf in distal cross
section, x300; 15. leaf cells at shoulder, xl25; 16. leaf apex, xl25; 17. leaf apex with gemmae, xl25; 18.
gemmae, x400. (1-9, Wager 125; 10-18, Crosby & Crosby 10136).
164
Calymperaceae
A genus of approximately 250 species, Calymperes is most common in tropical regions, especially in Africa
and southeast Asia. In Southern Africa the genus is very rare; the taxa were only recently reported from the
Flora area.
1 Teniolae present, occasionally weak above 1. C. rabenhorstii
1 Teniolae absent:
2 Costa excurrent; gemmae produced in cluster all around tip of costa 2. C. tenerum var. edamense
2 Costa percurrent; gemmae produced on ventral costal surface 3. C. levyanum
1. Calymperes rabenhorstii Hampe &
C. Mull, in Flora, Jena 69: 512 (1886);
Edwards in J. Bryol. 1 1 : 62 (1980). Type:
Guineae, Lagos, Rabenhorst s.n. (BM).
Plants small, in loose tufts, dark green;
corticolous. Stems erect, irregularly branched;
in section round, central strand absent, inner
cortical cells in 4-5 rows, outer cortical cells
in 1-2 rows, slightly smaller, brownish.
Leaves contorted and exposing hyaline bases
dry, erect-spreading wet; oblong to Ungulate
above obovate base, 2-3(4) mm long; apex
rounded, apiculate; base sheathing; margins
plane, serrate, strongly so at shoulders;
teniolae present, occasionally weak above, in
section forming intermarginal, multicellular
knob; extrateniolar lamina 1-3 cells wide.
Costa percurrent to just excurrent, ventral
superficial cells quadrate to rectangular,
smooth to weakly prorate above, dorsal
superficial cells rectangular, sharply prorate;
in section elliptical, stereids absent, guide
cells 6-8, large, weakly thickened, ventral
cells in 3-4 layers, smaller than guide cells,
ventral surface cells slightly larger, mam-
millose, dorsal cells in 6-7 layers, cells in 3
rows under guide cells similar to ventral
cells, 3-4 outer rows with cells smaller,
flattened, incrassate, dorsal surface cells
larger, sharply mammillose. Laminal cells
small, rounded-quadrate, ventral surface
strongly mammillose, dorsal surface weakly
mammillose or papillose; single row of
juxtacostal cells slightly larger; cancellinae
rectangular; teniolar cells rectangular to
long-rectangular, smooth.
Dioicous. Perichaetia terminal, leaves
weakly differentiated, base somewhat longer.
Seta 3-4 mm long, red-brown; capsule
cylindrical, 2,0-2, 5 mm long, yellow-brown;
exothecial cells rectangular, weakly sinuolate,
in zig-zaging rows, at mouth with 2 rows of
quadrate cells and up to 7 rows of transversely
rectangular cells below mouth; stomata
present at base of urn, phaneropore; peri-
stome absent; operculum long-rostrate,
0,7-1 ,2 mm long, cells not twisted; calyptra
persistent, cylindrical-rostrate, clasping base
of capsule, scabrous above, dehiscence of
spores through longitudinal slits in upper
calyptra; spores rounded, 15-25 jum, yellow-
ish, smooth with scattered large granules.
Fig. 45:9-16.
Endemic to Africa, C. rabenhorstii is most
commonly collected in west tropical Africa, however
specimens have also been reported from Zaire and
Angola. A single specimen, with sporophytes, is
known from Southern Africa. The specimen was
recently collected on Raphia palms at Kosi Bay, in
north-eastern Zululand. Map 60.
Voucher: Vahrmeijer PRE-CHI 2930.
The species can be separated from the other
Southern African taxa by the presence of teniolae;
although they are rather weak on the only collection
seen from the Flora area. The Southern African
specimen did not have gemmae, but they are reported
to occur in clusters at the leaf apex and are clavate
with a long tapering apex (Edwards, 1980).
2. Calymperes tenerum C. Mull. var.
edamense Fleisch., Musci FI. Buitenzorg 1:
275 (1904); Edwards in J. Bryol. 11: 83
(1980). Type: Java, Insel Edam, Musci
Frond. Archipelagi Indici, ser. II, no. 63
(MANCH).
Calymperes nashii Williams in Bull. Torrey bot.
Club 47: 391 (1920); Steere in Grout, Moss FI. N.
Amer. 1: 133 (1937). Type: Haiti, Port Margot,
Nash 51 (NY).
Plants small, dark green to yellow-green,
scattered or in small tufts; corticolous.
Stems erect, 1-2 mm high, unbranched; in
section without central strand, cortical cells
large, thickened, becoming smaller toward
margin. Leaves incurved or contorted dry,
erect-spreading wet; oblong or occasionally
weakly obovate, 2,0-2, 8 mm long; apex
acute to obtuse; base not broader than
lamina; margins plane, entire, thickened
above base, in section forming small knob
of 3-5 round cells. Costa short-excurrent,
with gemmiferous tip, dorsally spinulose
above base; in section oval, guide cells 6,
Calymperaceae
165
Fig. 45. — Calymperes tenerum var. edamense (1-8): 1. habit, xl; 2. habit, xlO; 3. leaf, X25; 4. leaf in
proximal cross section, x 170; 5. leaf in distal cross section, x 170; 6. junction of basal and laminal cells,
x 170; 7. upper laminal cells, x640; 8. gemmae, x95. C. rabenhorstii (9-16): 9. habit, xl; 10. habit, x 10; 11.
leaf, x25; 12. leaf in proximal cross section, x 170; 13. leaf in distal cross section, x 170; 14. cancellinae at
lower right margin, x 170; 15. junction of cancellinae and laminal cells at right leaf shoulder, showing teniola,
X 170; 16. cells at leaf apex, x 170. (1-8, Magill 5425; 9-16, Vahrmeijer PRE-CH 1 2930).
ventral substereid band 5 cells thick, cells
irregularly shaped, ventral surface cells
slightly smaller than laminal cells, sharply
mammillose, dorsal substereid band to 6 cells
thick, cells irregularly shaped, dorsal surface
cells round, sharply mammillose. Upper
laminal cells quadrate to hexagonal, mammil-
lose ventrally, weakly papillose dorsally;
marginal cells quadrate, smooth, rectangular
below; cancellinae rectangular; teniolae ab-
sent. Gemmae fusiform, in apical cluster on
all sides of excurrent costa.
Map 61. — • Calymperes tenerum var. edamense
X Calymperes levyanum
166
Calymperaceae
Sporophyte not known from Africa.
Fig. 45: 1-8.
New to Southern Africa, C. tenerum var. e da-
me rise has been collected in Java, west and east
Africa, the West Indies, Mexico and southern Florida,
U.S.A. The Southern African specimens were col-
lected on trees along Ku-Nkanini Stream in northern
Zululand and in coastal dune forest near Sodwana
Bay. Map 61.
Vouchers: Magill 5400, 5425.
This species is similar to C. victoriae Dix., but
differs in the excurrent costa, absence of teniolae and
smaller size. See additional notes on C. victoriae
under C. levyanum.
3. Calymperes levyanum Besch. in Annls
Sci. nat. Bot., ser. 8, 1: 273 (1896); Broth,
in Natiirl. PflFam. 10: 241 (1924); Flor-
schiitz, Mosses of Suriname 121 (1964). Type:
Nicaragua, Levy s.n. (NY, PC!).
Plants small, in loose tufts, dark green;
corticolous. Stems 4—6 mm high, occasionally
branched; in section round, central strand
absent, inner cortical cells in 5-6 rows,
large, slightly thickened, yellowish, outer
cortical cells in 1-2 rows, smaller, incrassate,
reddish. Leaves curled dry, wide-spreading
wet; linear, 5, 5-6,0 mm long; apex acute;
base elongate, to 1,5 mm long, abruptly
constricted to linear lamina; margins dentate
above, with multicellular teeth, serrate at
shoulders by projecting cells; in section
thickened above base, forming round or
quadrate knob, 4-5 cells thick, inner cells
incrassate, round. Costa percurrent, gemmi-
ferous at apex; in section wedge-shaped,
guide cells 4-8, indistinct, ventral cells in 2-3
layers, incrassate, dorsal stereid band 4-5
cells thick, dorsal surface cells substereids.
Upper laminal cells rounded, quadrate to
transversely short-rectangular, papillose with
4-6 low, blunt papillae over lumen; marginal
cells quadrate, slightly larger, smooth; basal
marginal cells rectangular; cancellinae quad-
rate, teniolae absent. Gemmae clavate, pro-
duced on ventral costal surface at apex.
Sporophyte not known in Africa. Fig. 44:
10-18.
New to Africa, C. levyanum was previously
known from Central America, West Indies and
northern South America. The new record was col-
lected on a rotting log in wet forest at Bloukrans
Pass in the southern Cape, at an elevation of 200 m.
Map 61.
Voucher: Crosby & Crosby 10136.
Calymperes victoriae Dix. from the Victoria
Falls, Zimbabwe, is similar but differs from this
species by its oblong leaves that abruptly constrict to a
short, spathulate, gemmiferous tip. The costa in
section has dorsal and ventral stereid bands and
incrassate surface cells. The leaves also have narrow
teniolae extending from the leaf base to the thickened
margin. Both species grow on wood or bark in moist
environments.
3. SYRRHOPODON
Syrrhopodon Schwaegr., Spec. Muse. Suppl. 2: 110 (1824); Broth, in Natiirl. PflFam. 10: 299
(1924); Sim, Bryo. S. Afr. 261 (1926). Lectotype species: S. gardneri (Hook.) Schwaegr., vide
Britt, in Britt., FI. Bermuda 436 (1916).
Plants dark green with conspicuous hyaline leaf bases, loosely caespitose or in small
tufts; corticolous, saxicolous or terricolous. Stems 2-20 mm long; central strand absent.
Leaves spirally twisted or curled dry, exposing bright hyaline sheathing bases; margins thick-
ened or with hyaline, cartilaginous border, strongly dentate to smooth. Laminal cells small,
papillose or sharply mammillose; cancellinae nearly filling base.
Perichaetia terminal. Seta erect; capsule immersed or exserted, ovoid to cylindrical;
peristome teeth 16 or absent; operculum rostrate; calyptra cucullate, smooth, deciduous.
Syrrhopodon is a genus of 270 species concentrated in the Tropics. The genus is well represented in the
Southern Hemisphere and extends northward into North America, but is absent from Europe and northern
Asia.
1 Leaves strongly toothed above, spinose at shoulders 1. S.gomesii
1 Leaves weakly serrate along hyaline border:
2 Leaf margins smooth below; margins of perichaetial leaves ± smooth 2. S. uncinifolius
2 Leaf margins frequently denticulate at shoulders; margins of perichaetial leaves sharply toothed at
shoulders 3. S. obliquirostris
Calymperaceae
167
1. Syrrhopodon gomesii P. Varde in
Svensk bot. Tidskr. 42: 253 (1948). Type:
Mozambique, Namuli Mts, Gomes & Sousa
3491, p.p. (PC!).
Plants medium-sized, dark green, scat-
tered or in small groups; corticolous. Stems
5-12 mm high; in section round, central
strand absent, inner cortical cells in 6-8 rows,
large, yellowish, outer cortical cells in 1-3
rows, stereids to substereids, reddish. Leaves
inrolled or curved dry, spreading wet; linear
above obovate base, 4, 5-5,0 mm long; apex
acute or rounded in gemmiferous leaves;
margins thickened above shoulders, irregu-
larly dentate with intermittent groups of
larger teeth, spinose at shoulders. Costa per-
current; in section subround, guide cells 4,
large, ventral stereid band 2-3 cells thick,
ventral surface cells large, incrassate, sharply
mammillose, dorsal stereid band 4-5 cells
thick, dorsal surface cells incrassate, sharply
mammillose. Upper laminal cells quadrate to
hexagonal, extending down leaf between
cancellinae and border, sharply papillose or
papillae sometimes bifid ; cancellinae quadrate
above, rectangular below; basal marginal
cells in 3-4 rows, linear, smooth.
Sporophyte not known. Fig. 46: 22-29.
New to Southern Africa, S. gomesii was described
from the Namuli Mountains of Mozambique (1537
AC). Two recent collections have been made on
wood in mountain forests of the northeastern Trans-
vaal. Map 62.
Vouchers: Crosby & Crosby 7552; Smook &
Phelan 865.
Syrrhopodon gomesii is distinct from other
Southern African species of the genus by its strongly
dentate and spinose leaves and less obvious hyaline
leaf base. The plants grow on tree trunks and wood
in dense indigenous forests.
2. Syrrhopodon uncinifolius C. Mull, in
Hedwigia 38: 96 (1899); Broth, in Natiirl.
PflFam. 10: 231 (1924); Sim, Bryo. S. Afr.
263 (1926); Dix. in Trans. R. Soc. S. Afr. 8:
189 (1920). Type: Cape, Montagu Pass,
Rehmann 129 (BOL!; PRE!).
Syrrhopodon erectifolius C. Miill. in Hedwigia 38:
96 (1899). Syntypes: Cape, Montagu Pass, Rehmann
128 (BOL!; NH!), 129 (NH!); Orange Free State,
Kadziberg, Rehmann s.n.
Plants small to medium, loosely caespi-
tose, dark green with conspicuous hyaline
leaf bases; terricolous, saxicolous or corti-
colous. Stems 5-10 mm high, unbranched; in
section round, central strand absent, cortical
cells large, in 8 rows, somewhat smaller
toward margin. Leaves spirally twisted or
tightly inrolled above sheathing, hyaline base
dry, erect-spreading wet; ligulate above
obovate base, 2, 5-4,0 mm long; apex acute;
margins plane to undulate, entire below,
serrate to toothed at apex, border of linear,
hyaline cells from base to just below apex,
cartilaginous. Costa percurrent to mucro-
nate; in section elliptical to wedge-shaped,
guide cells 4, incrassate, ventral stereid band
2 cells thick, infrequently substereids, ex-
posed, dorsal stereid band 2-3 cells thick,
dorsal surface cells undifferentiated, smooth.
Upper laminal cells quadrate, extending down
,r J 20
—
j—
f-
S-, ”• , 3,r y ,,
IBS
'll ^
1 1 1 lArifi10'
Brnn
r Tfrurrl
TulY
rO-hTT™
1 Jrrrrh4
znrni dqr
l+tjh
-Hnr
lYM44--UiP
tvrHrr
hTT 1 1
1 MUJ/-
>
TT\
JLLJlL 1 rn
5r| 1 1 i
Ml
A j rtff
t i
3r 1 1
(i/TTr
Httt
Jr
|| II || j — j — |
w-lV-'ri
10* 12*
18* IP
20*
22- 24* 26* 28*
3tr 32- 34* 36*
Map 62. — • Syrrhopodon obliquirostris
X Syrrhopodon gomesii
Map 63. — • Syrrhopodon uncinifolius
168
Calymperaceae
6
Calymperaceae
169
leaf between cancellinae and border, cells
with 4-6 low, blunt papillae; cancellinae
rectangular to quadrate. Gemmae clavate,
4-5 cells long, produced on ventral costal
surface at apex.
Perichaetia terminal; leaves ligulate
above long-sheathing base, margins smooth
throughout. Seta to 5 mm long, light brown ;
capsule short-cylindrical, to 1 mm long, light
brown; peristome fragile, teeth vertically
striate; operculum long-rostrate, beak ±
oblique; calyptra cucullate; spores round,
17-18 pm, granulose. Fig. 46: 13-21.
Endemic to Southern Africa, S. uncinifolius has
been collected in forests of the southwestern, southern
and eastern Cape, Natal, eastern Orange Free State,
Swaziland and the central, eastern and northern
Transvaal. Map 63.
Vouchers: Crosby & Crosby 8007; Kemp 1057;
Von Breitenbach 405.
This species is identified by its spirally twisted,
dark green upper leaf lamina and obvious, sheathing,
hyaline bases. It is not easily separated from S.
obliquirostris without fruiting material; see note
under that species.
3 Syrrhopodon obliquirostris C. Mull.,
Syn. Muse. 1: 543 (1849); Broth, in Natiirl.
PflFam. 10: 231 (1924); Sim, Bryo. S. Afr.
263 (1926); Dix. in Trans. R. Soc. S. Afr. 8:
189 (1920). Type: Cape, Grootvaderbosch,
Ecklon s.n.
Plants small, loosely caespitose, dark
green with conspicuous hyaline leaf bases;
corticolous. Stems 5-10 mm tall; in section
round, central strand absent, cortical cells
in 4-6 rows, large, thin-walled. Leaves
inrolled or curled dry, spreading wet;
Ungulate above obovate base, 2, 5-3,0 mm
long; apex broadly acute; margins plane,
entire to weakly serrate, toothed at apex,
occasionally a few teeth at shoulders;
bordered by elongate, hyaline cells, extending
from base to near apex, cartilaginous. Costa
percurrent, toothed dorsally at apex; in
section sub-elliptical, guide cells 4, incrassate,
ventral stereid band 2 cells thick, exposed,
dorsal stereid band 3-4 cells thick, dorsal
surface cells not differentiated, smooth. Upper
laminal cells quadrate, only slightly extended
down leaf between cancellinae and border,
cells with 4-6 low, blunt papillae; cancellinae
rectangular to quadrate. Gemmae clavate,
4-5 cells long, frequent, produced ventrally
at leaf apex.
Perichaetia terminal, leaves linear above
long, sheathing base; margins sharply toothed
at shoulders, frequently with smaller teeth
along upper margin. Seta to 4 mm long;
capsule short-cylindrical, 1,5 mm long;
peristome teeth linear, 0,1 mm long, whitish,
blunt apically, vertically striate; operculum
rostrate, beak oblique; calyptra cucullate;
spores round, 17-18 pm, granulate. Fig.
46: 1-12.
Endemic to Southern Africa, S. obliquirostris is
infrequently collected. The species occurs on bark or
rotting wood in forests of the southwestern and
southern Cape, Natal and eastern Transvaal. Map 62.
Vouchers: Sim 8728, 9122; Wager 288.
This species is very closely related to S. uncini-
folius and may only represent a form of that species.
A specimen that could be definitely considered type
material has not been located; however, the Wager
specimen cited above was identified by Dixon with
the indication ‘agrees well with specimens in the
British Museum Collection’, presumably the Ecklon
collection. The sharply toothed perichaetial leaves
separate specimens from S. uncinifolius and they
are provisionally placed here, although Muller
(1849) does not describe this character. The leaf
bases of S. obliquirostris are described as ‘ obsolete
denticulata', a character observed on the Wager
specimen and other collections.
Fig. 46. — Syrrhopodon obliquirostris (1-12): 1. habit, wet, xl; 2. habit, dry, xl; 3. habit, dry, X10; 4.
habit, wet, x 10; 5. leaves, x 12; 6. leaf in distal cross section, x 250; 7. leaf in proximal cross section, x 250;
8. right half of lamina at junction of cancellinae and laminal cells showing margin, x 120; 9. leaf apex, x 120;
10. left half of perichaetial leaf lamina at junction of cancellinae and laminal cells showing margin, x 120; 11.
perichaetial leaf apex, x 120; 12. part of capsule mouth with peristome teeth and spores, x 170. S. uncinifolius
(13-21): 13. habit, wet, x 1 ; 14. habit, dry, x 1 ; 15. habit, dry, x 10; 16. habit, wet, x 10; 17-18. leaves, X 12;
19. right side of lamina at junction of cancellinae and laminal cells showing margin, x 120; 20. upper laminal
cells, x 350; 21. leaf apex, x 120. S. gomesii (22-29): 22. habit, x 1 ; 23. habit, dry, x 10; 24. habit, wet, x 10;
25. leaves, x 12; 26. leaf in distal cross section, x 300; 27. right half of lamina at junction of cancellinae and
laminal cells showing margin, xl20; 28. upper laminal cells, x350; 29. leaf apex, xl20. (1-12, Wager 198;
13-21, Crosby & Crosby 8007; 22-29, Crosby & Crosby 7552).
170
Calymperaceae
4. HYPODONTIUM
Hypodontium C. Mull, in Hedwigia 38: 96 (1899); Broth, in Natiirl. PflFam. 10: 234 (1924).
Lectotype species: Hypodontium dregei (Hornsch.) C. Mull, (selected here).
Plants large, caespitose, glaucous-green to yellow-green; terricolous or saxicolous.
Stems erect, 20-30 mm tall; central strand present. Leaves incurled dry, spreading wet, linear
to lingulate above clasping base; margins unistratose, weakly bordered to mid-leaf or above,
involute above mid-leaf or only at apex. Costa strong, with dorsal and ventral stereid band,
ventral surface cells strongly differentiated.
Perichaetia terminal, leaves subulate, sheathing seta; capsule oval to short-cylindrical;
peristome teeth 16, triangular, occasionally with weak properistomal thickenings; operculum
rostrate; calyptra cucullate; spores large.
The genus Hypodontium contains two species, both known only from forests and woodlands of Zimbabwe
and the eastern and southern parts of the Flora area.
1 Leaf margins generally involute from mid-leaf to apex; dorsal junction of lamina and costa with large
spines above base; laminal cells with massive papillae; hyaline leaf border rarely reaching mid-leaf,
if so, then disappearing in involute margins 1 . H. dregei
1 Leaf margins plane or only involute at apex; dorsal costal surface smooth above, papillose below; leaf
cells with low, stout papillae, occasionally larger; hyaline leaf border extending to near apex
2. H. pomiforme
1. Hypodontium dregei {Hornsch.) C.
Mull, in Hedwigia 38: 97 (1899); Broth, in
Natiirl. PflFam. 10: 234 (1924). Syntypes:
Cape, Zwart-kei, Windvogelberg, Drege s.n.,
22 Nov. ; Table Mountain, Ecklon s.n. (BM !).
Syrrhopodon dregei Hornsch. in Linnaea 15: 116
(1841); Sim, Bryo. S. Afr. 263 (1926).
Syrrhopodon perichaetialis Bruch ex Krauss in
Flora, Jena 29: 132 (1846). Type: Cape, Uitenhage,
Winterhoek, Krauss s.n. (BM!).
Trichostomum cyathiforme Dix. in S. Afr. J. Sci.
18: 310 (1922). Hyophila cyathiforme (Dix.) Sim,
Bryo. S. Afr. 221 (1926). Type: Zimbabwe, Victoria
Falls, Sim 8934 (BM, holo.!; PRE!).
Plants medium to large, forming dense
cushions, glaucous-green to olive-green; saxi-
colous or corticolous. Stems 10-30 mm tall,
frequently weakly tomentose; in section
round, central strand small, weak, inner
cortical cells in 4-5 rows, large, thickened,
yellowish red, outer cortical cells in 1-2 rows,
substereids or incrassate, reddish. Leaves
contorted dry, erect-spreading wet; narrowly
lingulate above oval, hyaline base, 4-5 mm
long; apex acute; margins entire, involute
above mid-leaf, plane below, narrow border
of elongate, hyaline cells ending below
involute portion of margin or infrequently
extending into involute region, 2-6 cells wide
in base, 1-2 cells wide at mid-leaf. Costa
mucronate, ventral surface flat or convex; in
section subround, guide cells 4-6, large,
ventral stereid band 2-3 cells thick, occasion-
ally substereids, ventral surface cells large,
incrassate, strongly papillose, dorsal stereid
band strong, 5-6 cells thick, dorsal surface
cells incrassate, smooth, cells near junction of
costa and lamina strongly papillose or
spinose. Upper laminal cells quadrate, ventral
surface with long, massive, bifid papillae,
to 18 pm tall, dorsal surface with low, blunt
papillae; marginal cells in involute region
smooth; basal cells rectangular, hyaline,
weakly papillose in upper base, smooth
below.
Perichaetia terminal, leaves highly dif-
ferentiated, subulate above long-sheathing
base, 5-6 mm long, apices frequently reaching
capsule base. Seta erect, 5-6 mm long,
yellowish; capsule short-cylindrical, 1, 6-2,0
mm long, yellow-brown; peristome teeth 16,
triangular, 0,25 mm long, smooth, red-
yellow, frequently with weak properistomal
thickenings; operculum rostrate, 1 mm long;
calyptra cucullate, 3 mm long; spores round,
35-45 pm, irregularly granulate. Fig. 47:
10-17.
Hypodontium dregei is presently known from
Zimbabwe and Southern Africa. It is frequently
collected on rock, shallow soil over rock or on trees,
in forests and woodlands of the eastern and southern
Flora area. Map 64.
Calymperaceae
171
Fig. 47. — Hypodontium pomiformc (1-9): 1.
habit, wet, x 1 ; 2. habit, dry, x 1 ; 3. habit, x 10; 4.
leaves, X 14; 5. leaf in distal cross section, x 170; 6.
basal leaf cells (right half), xl20; 7. upper lamina
(right side), Xl20; 8. leaf apex, xl20; 9. part of
capsule mouth with peristome teeth and spores,
x60. H. dregei (10-17): 10. habit, wet, xl; 11.
habit, dry, xl; 12. habit, x 10; 13. leaves, x8;
14. leaf in distal cross section, x 170; 15. upper basal
cells between costa and margin, x 120; 16. upper
lamina (right half), xl20; 17. leaf apex, x 120.
(1-9, Crosby & Crosby 7738; 10-17, Crosby & Crosby
7496).
172
Calymperaceae
Vouchers: Brenan M2753; Cholnoky 640a;
Crosby & Crosby 7496; Magill 3525, 4314; Smook
1036; Thompson 3571; Von Breitenbach 137.
Vegetatively the species could be confused with
some pottiaceous genera, however the massive
ventral leaf cell papillae, and distinct border in the
lower leaf should separate this species. In addition,
when dry the plants have a very distinctive appearance
produced by the contorted upper lamina and con-
spicuous sheathing, hyaline bases.
2. Hypodontium pomiforme {Hook.) C.
MiXll. in Hedwigia 38: 97 (1899); Broth, in
Natiirl. PflFam. 10: 234 (1924). Type: Cape,
Swellendam, Burchell s.n. (BM, holo.!).
Weissia pomiformis Hook., Muse. Exot. 131 (1819).
Syrrhopodon pomiformis (Hook.) Hampe ex C. Mull.,
Syn. Muse. 1: 531 (1849); Sim, Bryo. S. Afr. 262
(1926).
Hypodontium pomiforme var. macowanianum C.
Mull, in Hedwiaia 38: 97 (1899). Tvpe: Cape, Bosch-
berg, MacOwan 9 (GRA!; MANCH!).
Plants medium to large, in cushions,
green to yellow-green; saxicolous or corti-
colous. Stems 20-50 mm tall, branched
below, frequently densely tomentose; in
section round, central strand small, inner
cortical cells in 5-6 rows, large, outer corti-
cal cells in 1-2 rows, small, incrassate, red-
dish. Leaves crisped above base dry, erect-
spreading wet, linear above obovate base,
3-4 mm long; apex acute; margins entire,
plane or involute at apex, narrow border of
elongate, hyaline cells extending from base to
near apex, rarely ending well below apex.
Costa percurrent, ventral surface strongly
convex; in section subround, guide cells 4-6,
ventral stereid or substereid band in 3-4
layers, ventral surface cells large, with
numerous small papillae, dorsal stereid band
3 cells thick, dorsal surface cells not dif-
ferentiated, proximally with massive, low,
blunt or bifid papillae, smooth distally.
Upper laminal cells rounded, quadrate, incras-
sate, ventrally with 2-4 low, C-shaped
papillae per lumen, occasionally papillae
larger, bifid, dorsally weakly papillose to
smooth; basal cells rectangular to elongate-
hexagonal, hyaline, smooth.
Perichaetia terminal, leaves highly dif-
ferentiated, linear-lanceolate above sheathing
base, 4-6 mm long. Seta to 8 mm long;
capsule oval, 2 mm long; peristome teeth 16,
triangular, smooth, reddish yellow; oper-
culum long-rostrate ; calyptra cucullate ; spores
round, 45-50 //m, tuberculate. Fig. 47: 1-9.
Endemic to Southern Africa, H. pomiforme is
infrequently collected in forests of the southwestern
and southern Cape. It has also been occasionally
collected in the eastern Cape, Transkei, Natal, Zulu-
land and eastern Transvaal. Map 65.
Vouchers: Boucher 3662; Crosby & Crosby 7738,
8170; Esterhuysen 20109; Taylor 9626.
Hypodontium pomiforme is a large, stout, yellow-
green plant growing on rock or rarely on bark, while
by comparison H. dregei is smaller, dark green to
glaucous-green and frequently collected on bark.
These characters, as well as those cited in the key,
will generally separate the species. However, all
these characters exhibit some variability and a few
specimens have been difficult to place. These speci-
mens can be identified using costal anatomy. The
costa of H. pomiforme, in cross section, is strongly
convex ventrally, while the costa of H. dregei is flat
to weakly convex ventrally but very prominent
dorsally.
173
ENCALYPTACEAE
Plants medium to large, forming large tufts, green to dark green; terricolous or saxi-
colous. Stems branched above, radiculose below; central strand present or absent. Leaves
little altered dry, larger above; oblong, elliptical or spathulate; apex broadly rounded to
short-acuminate, occasionally cucullate; margins plane or weakly revolute, entire or papillose.
Costa ending below apex to excurrent; in section with dorsal stereid band. Upper laminal cells
quadrate to hexagonal or irregular, slightly thickened, strongly papillose with large C- or O-
shaped papillae; basal cells fragile, rectangular, longitudinal walls thin, end wall thickened,
reddish; basal marginal cells narrow, thickened, forming distinct border.
Autoicous or dioicous. Perichaetia terminal on stems or branches, leaves frequently
smaller. Seta erect; capsule cylindrical, smooth or furrowed dry; exothecial cells rectangular,
quadrate at mouth; stomata scattered on lower urn; peristome single, double or absent,
teeth 16, lanceolate to linear, prostome occasionally present; operculum deciduous with
calyptra, long-rostrate; calyptra large, cylindrical-rostrate, smooth or scabrous, completely
covering capsule, base entire, erose, lacerate or fringed ; spores large, frequently ornate.
A family with two genera ; only one, Encalypta, occurs in Southern Africa.
ENCALYPTA
Encalypta Hedw., Spec. Muse. 60 (1801); Broth, in Natiirl. PflFam. 10: 241 (1924); Sim,
Bryo. S. Afr. 264 (1926); Flowers ex Grout, Moss FI. N. Amer. 1: 137(1938); Smith, Moss FI.
Brit. Irel. 206 (1978). Type species: not designated.
With characters of the family.
The genus Encalypta contains c. 35 species. The species are frequently widely distributed and found in
temperate to arctic or alpine regions. The plants are gametophytically similar to several genera of Pottiaceae
subfamily Pottioideae. The peristome, which can be single, double or absent, indicates that the family is a
transition group between the Haplolepidae and Diplolepidae, with a status similar to that of Orthotrichaceae
between the Acrocarpi and Pleurocarpi.
Two species are known from Southern Africa, both widespread in the Northern Hemisphere. Specimens
almost always have sporophytes in some stage of development. The large calyptra characterizes the genus.
1 Calyptra fringed at base; spores smooth to papillose; leaves short-acuminate, costa short-excurrent. .
1. E. ciliata
1 Calyptra entire to erose at base; spores warty distally; leaves acute to obtuse, somewhat cucullate, costa
ending below apex or percurrent 2. E. vulgaris
1. Encalypta ciliata Hedw., Spec. Muse.
61 (1801); Broth, in Natiirl. PflFam. 10: 242
(1924); Sim, Bryo. S. Afr. 264 (1926);
Flowers ex Grout, Moss FI. N. Amer. 1:
142 (1938); Smith, Moss FI. Brit. Irel. 207
(1978). Type: Europe.
Plants medium to large, forming cush-
ions, dark green; terricolous or saxicolous.
Stems 5-20 mm tall, branching above, radi-
culose below; in section round, central
strand medium-sized, inner cortical cells
large, in 5-6 rows, thin-walled, outer cortical
cells smaller, in 2 rows, incrassate, reddish.
Leaves crowded above, weakly contorted dry,
widespreading wet; lingulate to oblong or
elliptical, 3-6 mm long; apex short-acumi-
nate, base oblong; margins papillose, weakly
recurved for a short distance just above base,
plane distally. Costa very short-excurrent,
ventral superficial cells quadrate, papillose,
similar to laminal cells, dorsal superficial
cells linear, smooth below, sharply prorate
distally; in section subround, guide cells 4,
large, with several dorsal supplementary
cells, ventral cells in 2 rows, slightly thickened,
ventral surface cells smaller, papillose, dorsal
stereid band strong, in 2-4 rows, dorsal
174
Encalyptaceae
surface cells not differentiated, smooth.
Upper laminal cells hexagonal to subhexa-
gonal, slightly thickened, papillae 2-6, large,
C- or O-shaped; basal cells rectangular,
fragile, vertical walls thin, end walls thick-
ened, reddish; basal marginal cells linear,
forming distinct border 2-4 cells wide.
Autoicous. Perichaetia terminal, leaves
smaller, oval-acute, costa excurrent as short
awn. Seta erect, 6-10 mm long, red-brown;
capsules cylindrical, 2, 5-3,0 mm long,
yellowish; exothecial cells rectangular to
linear, strongly thickened; peristome fragile,
frequently missing on empty capsules, teeth
lanceolate, occasionally perforated, 125 pm
long, granulate; operculum long-rostrate,
1,6 mm long; calyptra cylindrical-rostrate,
4, 5-5,0 mm long, base with strongly dif-
ferentiated fringe of enlarged, rectangular
cells; spores discoid, 36-38 /tm, yellowish,
tetrad scar obvious, distal surface smooth,
granulate or occasionally with rounded
papillae. Fig. 48: 1-12.
Encalypta ciliata is known from North, Central
and South America, Europe, Asia and Africa. In
Southern Africa, the species is occasionally collected
on soil or rock in mountainous regions of the central
Cape, Natal and Lesotho. Map 66.
Vouchers: Hilliard & Burtt 10502; Magill 4412,
4697, 5899.
Although the fringe at the calyptra base is fragile,
as stated by Sim (1926), its detection was never a
problem. In addition, E. ciliata and E. vulgaris can be
separated by the shape of the leaf apex, costal length
and spore ornamentation.
The peristome of E. ciliata is very fragile and has
generally broken away by the time the capsule is
empty; thus older specimens could be confused with
E. vulgaris.
2. Encalypta vulgaris Hedw., Spec. Muse.
60 (1801); Broth, in Natiirl. PflFam. 10: 24
(1924); Flowers ex Grout, Moss FI. N. Amer.
1: 139 (1938); Scott & Stone, Moss. S.
Aust. 221 (1976); Smith, Moss FI. Brit. Irel.
207 (1978). Type: Europe.
Plants medium-sized, forming cushions,
green; saxicolous. Stems 5-20 mm, radicu-
lose below; in section round, central strand
present, inner cortical cells medium-sized,
thin-walled, in 5-6 rows, outer cortical cells
smaller, in 2 rows, reddish. Leaves erect with
lamina inrolled dry, widespreading wet;
lingulate to elliptical, 3-4 mm long; apex
acute to mucronate, frequently cucullate;
base oblong; margins plane, papillose. Costa
ending below apex or percurrent, ventral
superficial cells quadrate, papillose, similar to
laminal cells, dorsal superficial cells elongate,
smooth; in section guide cells 4, large, with
several dorsal supplementary cells, ventral
cells in 2-3 rows, large, weakly thickened,
ventral surface cells smaller, papillose, dorsal
stereid band strong, in 4-5 rows, dorsal
surface cells undifferentiated, smooth. Upper
laminal cells quadrate to hexagonal, weakly
thickened, papillae 1-4, large, C- or O-
shaped; basal cells highly differentiated,
fragile, rectangular, vertical walls thin, end
walls thickened, reddish; basal marginal cells
linear, thickened, forming distinct borders
3- 5 cells wide.
Autoicous. Perichaetia terminal, leaves
short, spathulate-obovate, apex obtuse. Seta
4- 6 mm long, red-brown; capsule cylindrical,
2, 5-3,0 mm long, yellow-brown, exothecial
cells rhomboidal to quadrate, thin-walled;
peristome absent; operculum long-rostrate,
to 2 mm high; calyptra cylindrical-rostrate,
5,0-5, 5 mm long, entire to erose at base;
spores with prominent tetrad scar, 34-37 pm,
distal surface warty, red-yellow. Fig. 48:
13-22.
Widespread in the Northern Hemisphere, E.
vulgaris is also known from Australia and Southern
Africa. In the Flora area, the species is found on soil
in rock crevices in mountainous areas of the south-
western and central Cape, Transkei and Lesotho.
Map 66.
Vouchers: Magill 4227; Schelpe 4927; Van Rooy
42.
Map 66. — • Encalypta ciliata
x Encalypta vulgaris
Encalyptaceae
175
Fig. 48. — Encalypta ciliata (1-12): 1. habit, dry,
X 1 ; 2. habit, wet, x 1 ; 3. habit showing a capsule
with calyptra and a deoperculate capsule, x 5 ; 4.
stem in cross section, X40; 5. leaf, xl2; 6. leaf in
cross section, x 143; 7. basal leaf cells at left margin,
xl25; 8. upper laminal cells, xl25; 9. leaf apex,
dorsal surface, xl25; 10. perichaetial leaf, xl2; 11.
perichaetial leaf apex, xl25; 12. part of capsule
mouth with peristome teeth and spores, xl60. E.
vulgaris (13-22): 13. habit, xl; 14. habit showing
deoperculate capsule, x5; 15. leaf, xl2; 16. leaf in
cross section, xl43; 17. basal juxtacostal cells,
xl25; 18. upper laminal cells, x500; 19. leaf apex,
xl25; 20. perichaetial leaf, xl2; 21. capsule with
calyptra, x5; 22. spores, x210. (1-12, Magill 4562;
13-22, Schelpe 4927).
176
Encalyptaceae
The erose to entire base of the calyptra is the
most obvious character to separate this species from
E. ciliata. In addition, the spores of E. vulgaris have
large wart-like processes on the distal face, while the
spores of E. ciliata are smooth to papillose distally.
The leaf apices of E. vulgaris in Southern Africa
are mostly obtuse or occasionally mucronate and
slightly cucullate, and the costa is percurrent to
subpercurrent. In the Northern Hemisphere, speci-
mens are known with piliferous apices and the costa
extending into the hair-point.
177
POTTIACEAE
Plants small to robust, usually in dense tufts; terricolous, saxicolous or corticolous.
Stems erect, sympodially branched or rarely monopodially branched ; central strand present
or absent. Leaves appressed or contorted dry, spreading to squarrose wet; linear to oblong or
spathulate, or ovate to lanceolate; lamina generally unistratose, rarely bistratose; apex acute to
broadly rounded; margins usually entire, occasionally crenulate to dentate above, plane
throughout to revolute or involute. Costa single, percurrent to excurrent as long or short
awn, smooth or denticulate, rarely ending below apex; in section ventral stereid band present
or absent. Upper laminal cells small, quadrate to hexagonal or rounded, mostly papillose,
occasionally mammillose or smooth, generally incrassate; basal cells generally larger, oblong,
smooth. Gemmae or propagulae occasionally produced on rhizoids, leaves or stem.
Perichaetia terminal or rarely lateral, leaves scarcely differentiated, rarely distinct. Seta
elongate, rarely shorter than urn; capsule symmetrical, stegocarpic, short- to long-cylindrical
or rarely cleistocarpic, globose to oval; peristome present or absent, single, teeth 16, erect to
spirally twisted, entire or divided into 32 filaments above membranaceous basal cylinder,
papillose to smooth ; operculum conical to rostrate ; calyptra cucullate ; spores mostly round,
smooth to strongly papillose.
Pottiaceae is by far the largest family of mosses with 89 genera. The family is also the largest in Southern
Africa, with 27 genera found most frequently in woodland, shrubland or grassland communities.
Key to Subfamilies and Tribes
1 Capsules cleistocarpic:
2 Costa in section reniform, ventral stereid band strong
Subfamily Trichostomoideae (p. 249)
2 Costa in section round, without ventral stereid band Tribe Pottieae (p. 192)
1 Capsules stegocarpic:
3 Costa without ventral stereid band, ventral cells occasionally incrassate
Subfamily Pottioideae (p. 178)
3 Costa with ventral stereid band strong or weak :
4 Stems without central strand :
5 Leaf margins recurved in base, frequently toothed above; laminal cells with
strongly spinose or forked papillae Tribe Leptodontieae (p. 186)
5 Leaf margins plane, entire; laminal cells with low, blunt papillae:
6 Plants small; peristome absent; leaves 1-2 mm long
Tribe Pleuroweisieae (p. 179)
6 Plants larger; peristome present; leaves 2, 5-5,0 mm long:
7 Leaves narrow, linear-lanceolate, lamina flat to weakly convolute
Subfamily Trichostomoideae (p. 249)
7 Leaves broad, ligulate, lamina rugose Tribe Barbuleae (p. 227)
4 Stems with central strand present but occasionally very weak :
8 Leaf margins completely or in part recurved to revolute. . . .Tribe Barbuleae (p. 227)
8 Leaf margins plane or involute:
9 Leaf margins involute
Subfamily Trichostomoideae (p. 249)
178
POTTIACEAE
9 Leaf margins plane:
10 Laminal cells smooth to mammillose:
11 Lamina bistratose juxtacostally Subfamily Trichostomoideae (p. 249)
11 Lamina unistratose Tribe Barbuleae (p. 227)
10 Laminal cells papillose:
12 Papillae C-shaped or rarely with 1-2 low, sharp, indistinct papillae per
cell; leaves ligulate to lanceolate, 1-2 mm long, or lamina broadly
oblong, 3-4 mm long, rugose Tribe Barbuleae (p. 227)
12 Papillae numerous, low and blunt, frequently obscuring cells, not C-
shaped; leaves linear to oblong or lanceolate, occasionally elliptical or
narrowly spathulate, 2-6 mm long . . Subfamily Trichostomoideae (p. 249)
Subfamily Pottioideae
Plants minute to large, gregarious or forming dense cushions; terricolous, saxicolous or
corticolous. Stems erect ; central strand present or absent. Leaves generally broad, orbicular or
oval to lanceolate or ligulate to spathulate; margins plane to revolute. Costa infrequently
bearing filaments or lamellae on ventral surface, percurrent to very long-excurrent, frequently
dentate; in section with or without ventral stereid band. Upper laminal cells large or small,
papillose or smooth; contact with basal cells straight across leaf or laminal cells extending
down margins forming distinct fl-shaped pattern at junction with basal cells. Gemmae fre-
quently produced on rhizoids, stems or leaves.
Capsules globose or oval to long-cylindrical ; peristome present or absent, teeth generally
long-filiform above basal membrane, erect to spirally twisted; operculum conic to rostrate;
calyptra cucullate.
Key to Tribes of Pottioideae
1 Ventral stereid band present, occasionally weak or absent in some leaves :
2 Stem with central strand:
3 Peristome absent or rudimentary; leaves narrow; margins plane or recurved on
one side Pleuroweisieae (p. 179)
3 Peristome well developed, if absent, leaves broadly elliptical ; margins revolute or
occasionally plane Barbuleae (p. 227)
2 Stems without central strand :
4 Plants large or wiry, very loosely caespitose; leaf margins recurved below, frequently
toothed above Leptodontieae (p. 186)
4 Plants small to medium, in dense cushions; leaf margins plane or recurved on one
side, infrequently toothed at shoulders Pleuroweisieae (p. 179)
1 Ventral stereid band absent:
5 Leaves mostly broadest at middle or above, orbicular to elliptical or Ungulate to
spathulate; leaf cells generally large with several C-shaped papillae, or infrequently
smooth or mammillose Pottieae (p. 192)
5 Leaves ovate to lanceolate, broadest below middle; papillae rarely C-shaped:
6 Leaf cells spinose-papillose; leaves in 3-ranks Leptodontieae (p. 186)
POTTIACEAE
179
6 Leaf cells with low, blunt papillae, smooth or mammillose ; leaves not 3-ranked :
7 Leaf cells with low, blunt papillae, frequently obscuring cells; leaf margins plane
Pleuroweisieae (p. 179)
7 Leaf cells mammillose, weakly papillose or smooth; leaf margins recurved to
revolute below, plane to revolute above or, if plane throughout, margins
bistratose Barbuleae (p. 227)
Tribe PLEUROWEISIEAE
Plants small to medium, in cushions; saxicolous or terricolous. Stems with or without
central strand. Leaves ligulate, lanceolate or triangular; apex obtuse to rounded or broadly
acute; margins frequently serrate to denticulate. Costa ending well below apex to percurrent;
ventral stereid band present or absent, dorsal stereid band strong. Upper laminal cells rounded,
quadrate to short-rectangular, papillae low, blunt, frequently obscuring cells; basal cells
moderately differentiated, rectangular, smooth.
Capsules terminal or on short lateral branches, stegocarpic; peristome absent or rudi-
mentary; operculum rostrate; calyptra cucullate; spores small.
Key to Genera of Tribe Pleuroweisieae
1 Stems monopodial; archegonia borne on short lateral branches; perichaetial leaves
strongly differentiated 1. Anoectangium
1 Stems sympodial; archegonia borne terminally on main stem; perichaetial leaves only
weakly differentiated :
2 Plants dark green, dull; leaves narrowly lanceolate; margins frequently denticulate at
shoulders, bistratose above 3. Gymnostomum
2 Plants light to yellowish green, if dark green then glossy; leaves ligulate to lanceolate;
margins entire, unistratose :
3 Plants often glossy; leaves strongly keeled; margins recurved on one or both sides
below; capsules systylious 2. Hymenostylium
3 Plants dull; leaves plane or weakly keeled; margins plane; operculum deciduous
3. Gymnostomum
1. ANOECTANGIUM
Anoectangium Schwaegr., Spec. Muse. Suppl. 1 : 33 (1811), nom. cons.; Saito in J. Hattori bot.
Lab. 39: 455 (1975). Type species: A. compactum Schwaegr.
Plants small, forming dense tufts. Stems monopodial; central strand present. Leaves
patent; ligulate to ovate-lanceolate. Costa with dorsal stereid band only. Laminal cells small,
obscure, papillose.
Perichaetial leaves distinct; capsule on short lateral branches, peristome absent; oper-
culum long-rostrate, deciduous.
There are c. 65 species of Anoectangium. The genus is found throughout the world, although the majority
of species are known from the Tropics. The plants occur most frequently on rock at upper elevations. In
Southern Africa, Anoectangium occurs throughout the Drakensberg range.
POTTIACEAE
181
Anoectangium wilmsianum (C. Mm//.)
Par., Ind. Bryol. Suppl. 13 (1900); Sim, Bryo.
S. Afr. 267 (1926). Type: Transvaal, Lyden-
burg, Wilms s.n. (G, holo. !).
Zygodon wilmsianum C. Mull, in Hedwigia 38:
113 (1899).
Anoectangium assimi/is Broth. & Wag. in Trans.
R. Soc. S. Afr. 4: 5 (1914). Type: Natal, Wager s.n.
(PRE!).
Plants slender, frequently very tall,
forming dense cushions, green to yellow-
green; saxicolous or infrequently terricolous.
Stems 5-20 mm tall, occasionally irregularly
branched below, sparsely tomentose below;
in section round to angular, central strand
small, rarely blackish, inner cortical cells lax,
outer cortical cells in 2 rows, stereids, red-
dish; axillary hairs of 7-8 cylindrical cells,
hyaline throughout. Leaves crowded, in-
curved, twisted dry, patent wet; ligulate to
lanceolate, high altitude specimens broader,
ovate-lanceolate, 0,8- 1,2 mm long; ventral
surface deeply and narrowly grooved along
costa; apex acute, mucronate by 1-3 smooth,
translucent cells; base scarcely differentiated,
frequently concave juxtacostally; margins
plane, entire. Costa strong, percurrent;
ventral superficial cells rectangular, smooth
or rarely papillose, dorsal superficial cells
long-rectangular, papillose; in section semi-
circular, lamina inserted ventrally, guide
cells 2, exposed, ventral cells generally
lacking, dorsal stereid band strong, 2-3 cells
thick, dorsal surface cells substereids. Upper
laminal cells rounded, quadrate to short-
rectangular or triangular, incrassate, papillae
low, blunt, scattered, 3-6 per cell ; basal leaf
cells quadrate marginally, papillose to near
base, juxtacostally quadrate to rectangular,
smooth.
Dioicous. Perigonia lateral, gemmate;
perichaetia on short lateral branches, leaves
very distinct, oval to ovate, abruptly acumi-
nate, 0,7-1 ,2 mm long, serrate at shoulders;
laminal cells rectangular, smooth. Seta 4-7
mm long, yellow; capsule oval, 1,0-1, 2 mm
long, reddish yellow; peristome absent;
operculum long-rostrate, 0,8 mm long;
calyptra cucullate; spores round, 12-13 pm,
spiculate. Fig. 49: 1-10.
Endemic to Southern Africa, A. wilmsianum
forms dense cushions on rock or soil over rock in the
Drakensberg of Lesotho, Natal, Orange Free State
and the eastern and northern Transvaal. Map 67.
Vouchers: Esterhuysen 26180; Magill 5762; Sim
9993; Symons 8662; Van Rooy 22.
The type specimen has the ligulate to lanceolate
leaves generally associated with A. wilmsianum. Recent
collections from high elevation in eastern Lesotho
and Giant’s Castle, Natal, have ovate-lanceolate
leaves, quite distinct from the type. The specimens
conform in other characters to A. wilmsianum. Varia-
tion in leaf shape expressed by all specimens examin-
ed, and an intermediate, described by Brotherus and
Wager (Wager, 1914) as A. assimilis, argue against
separation of the high altitude specimens at this time.
They may well require infraspecific status after further
study.
Fig. 49. — Anoectangium wilmsianum (1-10): 1. habit, xl; 2. habit, x6; 3. stem in cross section, x250;
4. leaves, x50; 5. leaf in cross section, x640; 6. cells at leaf base (papillae partly shown), x435; 7. upper
laminal cells at margin (papillae partly shown), x435; 8. leaf apex (papillae partly shown), x435; 9. perichaetial
leaf, x50; 10. operculum, x6. Hymenostylium recurvirostrum (11-21): 11. habit, Xl; 12. habit, x3; 13. stem
in cross section, x 250; 14. leaves, x 35; 15. leaf in proximal cross section, x 390; 16. leaf in distal cross section,
x 390; 17. lower left side of lamina (papillae partly shown), x 170; 18. upper laminal cells, x 640; 19. calyptra,
X20; 20. part of capsule mouth, x435; 21. systylious capsule, x 10. (1-2 & 9-10, Symons 8661 ; 3-8, Sim 9993;
11-21, Esterhuysen 26180).
182
POTTIACEAE
2. HYMEN OST YLIUM
Hymenostylium Brid., Bryol. Univ. 2: 81 (1827); Broth, in Natttrl. PflFam. 10: 257 (1924);
Gangulee, Moss. E. India 1 : 644 (1972). Type species: H. xanthocarpum (Hook.) Brid.
Plants in loose cushions, somewhat glossy; saxicolous. Stems sympodial, erect, to 20 mm
high; lacking central strand. Leaves strongly recurved wet, keeled, ligulate; apices acute.
Costa with dorsal and ventral stereid band. Lamina l cells quadrate to short-rectangular,
papillose.
Perichaetial leaves weakly differentiated; capsule systylious; peristome absent; oper-
culum long-rostrate, persistent.
The genus is known throughout the world, primarily through the wide distribution of H. recurvirostrum,
although 21 species have been described. Many authors treat these species under Gymnostomum. The systylious
capsule, with the operculum remaining attached to the columella and elevated above the capsule mouth when
dry, argues strongly for the recognition of Hymenostylium as a separate genus.
Hymenostylium recurvirostrum ( Hedw .)
Dix. in Revue bryol. lichen. 6; 96 (1934);
Zander in Bryologist 80: 253 (1977). Type:
Europe, Ehrhart.
Gymnostomum recurvirostre Hedw., Spec. Muse. 33
(1801); Saito in J. Hattori bot. Lab. 39: 452 (1975).
Plants small, loosely caespitose, yellow-
green to dark green, frequently glossy;
saxicolous. Stems 10-20 mm high, frequently
branching by subperichaetial innovations,
occasionally papillose, frequently with red-
dish tomentum below; in section round to
angular, central strand absent, inner cortical
cells large, thin-walled, outer cortical cells in
1-2 rows, smaller, incrassate, reddish; axil-
lary hairs cylindrical, 6-9 cells long, hyaline
throughout. Leaves somewhat distant, twisted
and appressed dry, recurved to squarrose wet,
generally keeled; ligulate to oblong-lanceo-
late, (0,5-) 1-2 mm long; apex acute; base
scarcely differentiated to oval; margins
entire, plane to recurved on one or both sides
in lower leaf. Costa percurrent to just
excurrent ; ventral superficial cells rectangular,
smooth, dorsal cells long-rectangular, papil-
lose, rarely smooth; in section semicircular to
oval, guide cells 2, ventral stereid band weak,
1 (-2) layer(s) thick, ventral surface cells
rarely present, thin-walled, dorsal stereid
band 3-4 layers thick, dorsal surface cells
distinct, outer wall strongly thickened, lumen
crescent-shaped. Upper laminal cells quadrate
to short-rectangular, irregularly thickened,
especially at corners, papillae scattered, low,
blunt, 2-3 per cell; basal cells moderately
differentiated, rectangular, smooth, thin-
walled.
Dioicous. Perigonia terminal, gemmi-
form; perichaetia terminal, leaves weakly
differentiated, 1 , 5 mm long. Seta 4-8 mm
long, reddish brown to yellowish; capsule
systylious, urn oval to short-cylindrical,
0,8-1, 2 mm long, reddish brown; peristome
absent; operculum obliquely rostrate, 0,4-
0,8 mm long, cells not twisted; calyptra
cucullate, 1,2-1, 5 mm long; spores round,
10-12 pm, granulate, brown. Fig. 49: 11-21.
This species is known from North, Central and
South America, Europe, Africa, Asia and Australasia.
In Southern Africa, specimens of H. recurvirostrum
are collected on damp or moist rock, generally in
,r
1* , f ,
7* , J(r ,jf
J 1 ITTfl
.r
IT — l — l-
-1 N — P
Y
H — i
HH
TTT
ILL TTl
,0*
TM 1 r
—
-U — L
Till
ff-
ir
p — fpT I |
-mTm
P
1 \l I 1 j / 1
»•
— r
TTxru
1 1
IL
.
~N
j
TT 7
+44-firr
IT \
TP
-
-
i
~T
ir
Htttv
_
-HTnT
ill
T
,0*
-4-
\
—
•
jy| 1 1 1 rn”’
jT
T44-
TP
k
-
_
yy ]y
iTTTTrn
-4-
y=
Httt
TkTT
P
n+LM-fl
44+rr
' Jp ir
1 Jr
_
_J
rr
1 1 1 j j j /I
Map 68. — • Hymenostylium recurvirostrum
POTTIACEAE
183
association with waterfalls or seepage areas. Most
Southern African specimens have come from the
Drakensberg of western Natal and Lesotho; however,
a few collections have been made in shaded creek beds
or rock cliffs in central South West Africa/Namibia,
central Cape and eastern Transvaal. Map 68.
Vouchers: Esterhuysen 21624; Magill 4833;
Phelan & Smook 73 ; Schmitz 8101.
Hymenostylium recurvirostrum is identified by its
glossy appearance, keeled leaves, and lower leaf
margins generally recurved on one side. When sporo-
phytes are present, the persistent operculum, even on
empty capsules, will separate the species from other
taxa in the Flora.
The east African species, H. crassinervium, is
very similar, but differs in linear to narrowly ligulate
leaves, short-excurrent costa and a single low, blunt
papilla per lumen. The species is known as far south
as Zimbabwe, but has not been recorded from South-
ern Africa.
3. GYMNOSTOMUM
Gymnostomum Nees & Hornsch., Bryol. Germ. 1 : 153 (1823), nom. cons.; Saito in J. Hattori
bot. Lab. 39: 450 (1975). Type species: G. calcareum Nees & Hornsch.
Plants small to medium, dark green to glaucous-green; terricolous or saxicolous. Stems
sympodial; central strand present or absent. Leaves ligulate to linear-lanceolate, unistratose
or bistratose; apex obtuse to acute or frequently rounded; margins entire to denticulate at
shoulders. Costa percurrent to ending below apex. Laminal cells quadrate, papillose.
Perichaetia terminal, leaves only weakly differentiated; peristome absent; operculum
rostrate, deciduous.
The 14 species of Gymnostomum are widely distributed throughout the world. The variable species G.
aeruginosum is almost cosmopolitan and generally found in association with calcareous or other basic rock.
Gymnostomum lingulatum and G. bewsii are endemic to Southern Africa.
1 Central strand present; leaves ligulate to linear-lanceolate, 0,4-1, 5 mm long 1. G. aeruginosum
1 Central strand absent; leaves 1-2 mm long:
2 Leaves lanceolate to ± triangular, unistratose; margins entire 2.G. lingulatum
2 Leaves linear to linear-lanceolate, bistratose; margins frequently denticulate at shoulders 3. G. bewsii
1. Gymnostomum aeruginosum J. E. Sm.,
FI. Brit. 3: 1163 (1804); Zander in Bryologist
80 : 259 ( 1 977). Type : Europe.
Gymnostomum calcareum Nees & Hornsch., Bryol.
Germ. 1 : 53 (1823). Type: Germany.
Plants small, in loose tufts or cushions,
yellow-green to light green, brownish below;
terricolous or saxicolous. Stem 1,0-2, 5 mm
tall, simple or sparsely branched below; in
section angular, central strand small, inner
cortical cells lax, outer cortical cells in 1-2
rows, incrassate, reddish; axillary hairs 3-6
cells long, hyaline throughout. Leaves incur-
ved to appressed dry, recurved wet; ligulate
to linear-lanceolate, 0,4- 1,5 mm long;
apex obtuse to rounded or broadly acute and
apiculate; base oblong; margin plane, entire.
Costa ending 2-5 cells below apex to per-
current; ventral superficial cells quadrate to
rectangular, sparsely papillose, occasionally
densely papillose above, dorsal superficial
cells rectangular, papillose; in section semi-
circular, guide cells 2, ventral stereid band
absent or rarely 1-2 cells over guide cells,
ventral surface cells similar to guide cells,
papillose, dorsal stereid band strong, 4 cells
thick, dorsal surface cells rarely differentiated,
papillose. Upper laminal cells quadrate or a
few short-rectangular, weakly thickened,
papillae crowded, low, simple, 4-6 per cell;
basal cells hyaline, rectangular, thin-walled,
smooth.
Dioicous. Perigonia gemmate; peri-
chaetia terminal, leaves weakly differentiated,
ovate-lanceolate, sheathing below, 1 , 5 mm
long. Seta 4-5 mm long, yellowish; capsule
elliptical, 0,5-0, 8 (-1) mm long, yellow-red;
peristome absent; operculum rostrate, to 0,5
mm long; calyptra cucullate, 1,2 mm long;
spores round, 9-12 pm, granulate. Fig. 50:
18-26.
184
POTTIACEAE
Gymnostomum aeruginosum is known from North,
Central and South America, Europe, Africa and
Asia. In Southern Africa the species is collected on
sandstone or other basic rocks or soils. It is rare in
the central and southern Cape and South West
Africa/Namibia and infrequent in the Drakensberg
of Natal, Lesotho and Orange Free State. Map 69.
Vouchers: Cholnoky 1041; Gibb PRE-CH6037;
Magill 4278; Schmitz 8106; Sim 9057.
Specimens of G. aeruginosum are recognized by
their small size, ligulate leaves with obtuse apices and
the costa not reaching the apex. The low, simple
leaf cell papillae should indicate the relationship to
Pleuroweisieae, and the terminal sporophytes and
early deciduous operculum will separate specimens
from other related genera.
2. Gymnostomum lingulatum Rehm. ex
Sim, Bryo. S. Afr. 260 (1926). Type: Trans-
vaal, Lechlaba, Rehmann 437 (PRE, holo.!;
BM!; BOL!).
Didymodon lingulatum (Sim) Magill in Mem. bot.
Surv. S. Afr. 43:5 (1979).
Plants medium-sized, loosely caespitose,
dark green above, reddish brown below;
?saxicolous. Stems 20-30 mm tall, branching,
mainly above, sparsely red-tomentose below;
in section subquadrate, central strand absent,
inner cortical cells lax, outer cortical cells in
1-2 rows, incrassate, reddish; axillary hairs of
6 cylindrical cells, hyaline throughout or
basal cell shorter, brownish. Leaves some-
what crowded, incurved, weakly twisted dry,
patent wet; lanceolate to triangular, 1,0- 1,5
mm long; concave ventrally; apex obtuse to
acute; base scarcely differentiated; margins
plane, entire. Costa subpercurrent to ending
4-5 cells below apex; ventral superficial cells
rectangular, smooth, dorsal superficial cells
rectangular, papillose; in section semi-
circular, guide cells 4-6, ventral substereid
band weak, of 1-2 cells, ventral surface cells
incrassate or substereids, dorsal stereid band
1-2 rows thick, dorsal surface cells sub-
stereids. Upper laminal cells rounded,
quadrate or occasionally a few cells short-
rectangular or triangular, incrassate, papillae
low, simple, 2-4 scattered over cell ; basal cells
weakly differentiated, rectangular to short-
rectangular, hyaline, smooth, thin-walled.
Sporophyte unknown. Fig. 50: 10-17.
Endemic to Southern Africa, G. lingulatum is
only known from the type locality in the northern
Transvaal. Map 70.
Voucher: Type only.
Re-examination of several plants indicated that
the basal cells of axillary hairs may be either hyaline
and undifferentiated or brownish and smaller. The
variation found by Zander (1977) in axillary hairs of
Gymnostomum, together with the absence of a central
strand in the stem and the low, simple, scattered leaf
cell papillae of G. lingulatum , indicate that the species
may be more correctly placed in Gymnostomum.
The plants examined have lanceolate or occa-
sionally triangular leaves. The specific epithet lingu-
latum, used by Rehmann on the specimens of his
exsiccate and later adopted by Sim (1926), may
indicate a greater variation in leaf shape on other
duplicates.
3. Gymnostomum bewsii Dix. in Trans. R.
Soc. S. Afr. 8: 190 (1920); Sim, Bryo. S. Afr.
257 (1926). Type: Transkei, Drakensberg,
Ongeluks Nek, Rowlins sub Sim 8251 (BM,
lecto. !, selected here; PRE!).
Fig. 50. — Gymnostomum bewsii (1-9): 1. habit, xl; 2. habit, xlO; 3. stem in cross section, x225; 4.
leaves, x40; 5. leaf in proximal cross section, x640; 6. leaf in distal cross section, x640; 7. laminal cells at
leaf shoulder, xl70; 8. leaf apex, xl70; 9. laminal cells at perichaetial leaf shoulder, x 170. G. lingulatum
(10 17): 10. habit, x 1 ; 11. habit, x 5; 12. stem in cross section, x 225 ; 13. leaves, x 40; 14. leaf in cross section
x 640; 15. upper laminal cells, x640; 16. basal leaf cells, x 170; 17. leaf apex, x 170. G. aeruginosum (18-26):
18. habit, x 1 ; 19. habit, x 10; 20. stem in cross section, x 320; 21. leaves, x 40; 22. leaf in cross section, x640;
23. basal leaf cells, x435; 24. upper laminal cells (papillae partly shown), x640; 25. leaf apex, x435; 26.
capsule, X 20. (1-2, Magill 5745; 3-9, Wager 159; 10-17, Rehmann 437; 18-26, Wager 473).
POTTIACEAE
185
186
POTTIACEAE
Plants slender, frequently very tall,
forming tufts or dense cushions, dark green to
light or yellowish green above, light brown
below; terricolous or saxicolous. Stems erect,
5-20 mm tall, frequently branching; in section
round to oval, central strand absent, inner
cortical cells lax, outer cortical cells in 1-3
rows, somewhat smaller, incrassate, reddish;
axillary hairs 5-10 cells long, hyaline through-
out. Leaves incurved, slightly twisted dry,
widespreading wet; linear to linear-lanceo-
late, 1-2 mm long; broadly concave ventrally,
bistratose above base; apex acute; base
scarcely differentiated, oblong to oval, erect-
appressed wet ; margins plane, entire or
infrequently denticulate at shoulders. Costa
strong, percurrent or just excurrent; ventral
superficial cells quadrate to short-rectangular,
weakly papillose, dorsal superficial cells
rectangular, smooth; in section elliptical to
wedge-shaped, guide cells 4-6, large, ventral
stereid band 1-2 cells thick, ventral surface
cells incrassate, dorsal stereid band 2-3 cells
thick, dorsal surface cells with strongly
thickened outer walls, lumen semicircular.
Upper laminal cells quadrate, frequently
transversely rectangular at margins, incras-
sate, papillae low, blunt, simple, 2-4 over
lumen; basal cells moderately differentiated,
bulging, quadrate to short-rectangular,
hyaline, smooth, thin-walled.
Dioicous. Perichaetia terminal, leaves
lanceolate to linear, 1,5 mm long, generally
dentate to denticulate at shoulders. Seta
2, 5-3,0 mm long, yellowish; capsule very
shortly cylindrical, 1 ,0-1 ,2 mm long, reddish
yellow; peristome absent; operculum long-
rostrate, 0,8 mm long, cells not twisted;
spores round, 12-17 pm, papillose, brownish.
Fig. 50: 1-9.
Endemic to Southern Africa, G. bewsii is infre-
quently collected in the Drakensberg of Natal,
Lesotho, Transkei and Orange Free State, and rarely
in the eastern Transvaal. Map 70.
Vouchers: Esterhuysen 34595; Hilliard & Burtt
10411; Magill 4719, 5745.
Map 70. — • Gymnostomum bewsii
x Gymnostomum lingulatum
A few of the specimens cited by Dixon (1920)
were misidentified and have been referred to Anoec-
tangium wilmsianum (cf. Magill & Schelpe, 1979). The
large, fruiting specimen, Rowlins sub Sim 8251 (BM!),
is selected as lectotype, because it most closely
matches the protologue. The specimen was cited by
Dixon (1920) as ‘Coll. Bro. Mayol, comm. Sim (No.
8251)’, but original correspondence and specimens at
PRE, indicate that the specimen was collected by Mr
Rowlins, sent by Bro. Mayol to Sim, who sub-
sequently sent a specimen to Dixon.
The dark green, dense cushions, with linear,
bistratose leaves will help to identify collections. The
infrequently denticulate margins at the leaf shoulders,
especially prominent in perichaetial leaves, were
responsible for the erroneous reports of Eucladium in
Southern Africa. The specimens of E. verticillatum
cited by Sim (1926) are G. bewsii. The accurate illus-
tration of E. verticillatum published by Sim (1926)
was apparently based on European plants; see com-
ment on his figure.
Tribe LEPTODONTIEAE
Plants medium to large, forming loose tufts or mats; terricolous or saxicolous. Stems
without central strand, round or triangular, frequently fluted. Leaves ovate to ovate-lanceolate;
acute to acuminate; margins recurved below, dentate or entire above. Costa generally with
dorsal and ventral stereid bands. Laminal cells rounded, quadrate to hexagonal, incrassate,
corners frequently thickened, spinose-papillose or with numerous low papillae over lumen,
occasionally forming crown-shaped group.
POTTIACEAE
187
Sporophyte terminal, perichaetial leaves differentiated. Capsule stegocarpic, cylindrical;
peristome present, short, erect; operculum rostrate.
Key to Genera of Tribe Leptodontieae
1 Leaves short, to 2 mm long, ovate to broadly ovate; basal leaf cells not differentiated
1 . Triquetrella
1 Leaves longer, 2, 5-5,0 mm long, ovate-lanceolate; basal leaf cells differentiated. . . .
2. Leptodontium
1. TRIQUETRELLA
Triquetrella C. Mull, in Ost. bot. Z. 47: 421 (1897); Broth, in Naturl. PflFam. 10: 264 (1924);
Sim, Bryo. S. Afr. 251 (1926). Lectotype species: T. tristicha (C. Mull.) C. Mull., vide Grout,
Moss FI. N. Amer. 1 : 170 (1938).
Plants wiry, forming loose cushions or mats. Stems triangular in section. Leaves 3-ranked,
ovate, short-acuminate; leaf base decurrent. Laminal cells quadrate to rhomboid, spinose-
papillose; basal leaf cells not differentiated. Sporophyte not known from Southern Africa,
similar to Leptodontium.
Of the 1 1 species of Triquetrella, only three are kn
are found in Australia, New Zealand, South America,
of these species appear to be very closely related and
tentative.
Triquetrella tristicha (C. Mull.) C. Mull.
in Ost. bot. Z. 47: 422 (1897); Broth, in
Naturl. PflFam. 10: 265 (1924); Sim, Bryo. S.
Afr. 251 (1926). Type: Cape, Swartkop River,
Ecklon s.n. (BM!).
Zygodon tristichus C. Miill. in Bot. Ztg 13: 764
(1855).
Triquetrella strictissima Rehm. ex C. Miill. in
Ost. bot. Z. 47: 422 (1897). Type: Cape, Wellington,
Rehmann 144 (BM!; PRE!).
Plants medium-sized, slender, in loose
cushions or mats, yellow-green to dark green,
brownish below; terricolous or saxicolous.
Stems erect or inclined, to 25 mm long,
rarely branched; in section triangular, central
strand absent, inner cortical cells large,
incrassate, yellowish, outer cortical cells in
1-2 rows, stereids, reddish yellow; axillary
hairs 6-8 cells long, cylindrical, 1-2 basal
cells very short, hyaline throughout. Leaves
distant below, crowded above, 3-ranked,
appressed, twisted dry, patent wet; ovate to
broadly ovate or oval, 1, 2-2,0 mm long;
apex acuminate; base decurrent; margins
recurved to mid-leaf, entire. Costa percurrent
to subpercurrent; ventral superficial cells
rectangular, smooth, dorsal superficial cells
own from the Northern Hemisphere. The other species
Southern Africa, and the Subantarctic Islands. Several
the present separation of many of the species is very
rectangular to quadrate above, papillose;
in distal section round to subround, frequent-
ly undifferentiated, especially in upper leaf,
cells irregularly placed, incrassate; in proxi-
mal section guide cells 2, large, incrassate,
ventral stereid band of 1-2 cells, surface cells
undifferentiated, dorsal stereid band weak,
1-2 rows thick, dorsal surface cells sub-
stereids, outer wall strongly incrassate.
Upper laminal cells quadrate to short-rhombic,
incrassate, in regular rows, papillae strong,
spinose or bifid, mostly 1 per cell; basal cells
undifferentiated or with small group of
rectangular, smooth cells at insertion near
costa.
?Dioicous. Perigonia gemmate, leaves
sheathing, oval-acuminate, laminal cells
rhomboid, smooth. Other parts not seen.
Fig. 51:1-8.
Triquetrella tristicha is common in the southern
Cape Province and has also been collected in the
eastern Orange Free State and western Lesotho.
Most specimens are collected in semi-arid regions;
only a few collections have been made in mesic
environments. A specimen from Kaapsche Hoop,
Transvaal ( Wager 644, PRE) appears oddly dis-
placed; other Transvaal collections have not been
seen. Map 71.
188
POTTIACEAE
Map 71. — • Triquetrella tristicha
x Leptodontium viticulosoides
Vouchers: Barnard 49298; Esterhuysen 16598;
Jacot Guillarmod 6324; Magill 6341; Magill &
Schelpe 3941 ; Wager PRE-CH7675.
Specimens of T. tristicha have not been collected
in Southern Africa with sporophytes, although a
recent collection ( Bayliss 8522) contained plants with
fully developed antheridia. A very closely related
Australian species, T. papillata (Hook. f. & Wils.)
Broth., is said to have ‘narrowly cylindrical capsules
on a long flexuous seta’ and was illustrated (Scott &
Stone, 1976) with long-sheathing perichaetial leaves.
Fro. 51. — Triquetrella tristicha: 1. habit, xl;
2. habit, x 10; 3. stem in cross section, xl50; 4.
leaves, x40; 5. leaf in cross section, x340; 6. cells
at leaf base (right side), x 170; 7. upper laminal cells,
x640; 8. leaf apex, xl70. (1-8, Magill 3866).
POTTIACEAE
189
2. LEPTODONTIUM
Leptodontium (C. Mull.) Hampe ex Lindb. in Ofvers. Forh. Kongl. Svenska Vetensk.-Akad.
21: 227 (1864); Zander in Bryologist 75: 230 (1972). Type species: L. squarrosum (Hook.)
Hampe ex Lindb.
Plants large, forming loose tufts, erect. Stems round, fluted or smooth in section.
Leaves many-ranked, ovate-lanceolate; base not decurrent. Laminal cells rounded, quadrate to
hexagonal, corners generally thickened, papillae scattered over lumen, numerous, or forming
massive crown-shaped group; basal cells rectangular, smooth, thin-walled. Gemmae on stem
obovoid, multicellular.
Sporophyte terminal; seta elongate; capsule cylindrical; peristome erect, teeth linear,
short; operculum rostrate; calyptra cucullate, large; spores papillose.
The genus Leptodontium presently comprises about 80 species that are widely distributed in tropical and
subtropical regions. In Southern Africa the genus is found along the Drakensberg escarpment of Transvaal,
Natal and Lesotho.
1 Stem smooth, not fluted in section 1. L. viticulosoides
1 Stem fluted in section:
2 Leaves 3-5 mm long; leaf cells incrassate, corners strongly thickened, in section cells bulging with
massive crown-shaped papillae 2. L. longieaule
2 Leaves 2,0-2, 5 mm long; leaf cells ± thin-walled, corners not thickened, in section cells ± flat,
papillae numerous, scattered over lumen 3. L. brachyphyllum
1 . Leptodontium viticulosoides ( P . Beauv.)
Wijk & Marg. in Taxon 9: 51 (1960); Zander
in Bryologist 75: 243 (1972). Type: Reunion,
Bory de St Vincent s.n.
Neckera viticulosoides P. Beauv., Prodr. 78 (1805).
Didymodon squarrosus Hook., Musci Exot. 2: 150
(1819). Trichostomum squarrosum (Hook.) Schwaegr.,
Spec. Muse. Suppl. 2: 78 (1823). Leptodontium
squarrosum (Hook.) Hampe ex Lindb. in Cjfvers.
Forh. Kongl. Svenska Vetensk.-Akad. 21 : 227 (1864);
Sim, Bryo. S. Afr. 249 (1926). Type: Nepal, Gardner
s.n. (BM, holo. !).
Zygodon simii Dix. in Trans. R. Soc. S. Afr. 8:
198 (1920). Type: Natal, Swartkop, Sim 8690 (BM,
holo.!; PRE!).
Plants large, in loose tufts, green to
yellow-green, brown below; terricolous, saxi-
colous or corticolous. Stems 20-80 mm tall,
frequently covered with white tomentum,
reddish below; in section round, not fluted,
central strand absent, inner cortex large,
cells thin-walled, outer 1-2 rows stereids,
reddish; axillary hairs 15-16 cells long,
filamentous, hyaline throughout. Leaves
crowded, erect-spreading, twisted or con-
torted dry, squarrose to recurved wet,
weakly keeled; ovate-lanceolate to broadly
lanceolate, 3,0-3, 5 mm long; apex acuminate,
occasionally acute; base ovate, sheathing;
margins narrowly recurved below, plane
above, dentate in upper leaf. Costa sub-
percurrent; ventral superficial cells rectangu-
lar, smooth, dorsal superficial cells rectangu-
lar, weakly papillose; in section reniform,
guide cells 2-4, large, ventral stereid band
1 (-2) layers thick, exposed, dorsal stereid
band 2-3 layers thick, surface cells not
differentiated. Upper laminal cells rounded-
quadrate, corners strongly thickened, with
2-4 small, simple papillae scattered over the
cell; basal cells moderately differentiated,
long-rectangular, slightly thickened, porose.
Autoicous. Perigonia lateral; perichaetia
terminal; leaves long-sheathing, 9-10 mm
long. Seta 12-15 mm long, yellowish; capsule
cylindrical, 4 mm long, red-yellow; peristome
short, 0,1 mm long, teeth linear, erect,
irregular, properistome present; operculum
rostrate, 1,5 mm long; calyptra 6 mm long;
spores round, 22-25 pm, weakly papillose.
Fig. 52: 1-9.
Specimens are frequently collected, from a large
variety of substrates, in the forests of the northern
and eastern Transvaal and Natal. Map 71.
Vouchers: Magill 3404; Sim 8705; Smook 848;
Von Breitenbach 139; Wager 121.
The large plants with squarrose-recurved, den-
tate leaves and white tomentum will help to identify
L. viticulosoides. The species can be separated from
the other two by its non-fluted stems and low, simple
leaf papillae.
190
POTTIACEAE
19
POTTIACEAE
191
2. Leptodontium longicaule Mitt, in J.
Linn. Soc., Bot. 12: 51 (1869); Zander in
Bryologist 75: 266 (1972). Type: Ecuador,
Pichincha, Spruce 306 (NY, holo. ; BM !).
Plants medium to large, forming large
tufts, green to yellow-green, brownish below;
terricolous or saxicolous. Stems 10-80 mm
tall, rarely branched, without tomentum; in
section oval, fluted, central strand absent,
cells of cortex large, incrassate, strongly
thickened toward margin, outer 1-3 rows
stereids, reddish; axillary hairs of 14-17 short
cells, basal cell brownish. Leaves crowded,
twisted to contorted above with appressed
bases dry, strongly squarrose-recurved wet;
ovate-lanceolate, 3-5 mm long; apex acute;
base oblong, sheathing; margins narrowly
recurved below, serrate in upper leaf. Costa
subpercurrent ; ventral superficial cells long-
rectangular, smooth, dorsal superficial cells
long-rectangular, finely papillose; in section
reniform, guide cells 2-4, large, ventral
stereid band 1-2 cells thick, exposed, dorsal
stcreid band 2-3 cells thick, dorsal surface
cells not differentiated. Upper laminal cells
rounded-quadrate, thickened at corners,
bulging superficially, papillae crown-like,
massive, centered over lumen; basal cells
moderately differentiated, rectangular, thick-
ened, pitted, seriate papillose to lower base.
Sporophyte not known in Southern
Africa. Fig. 52: 10-16.
Leptodontium longicaule is known from Central
and northern South America, eastern and Southern
Africa and several African and American islands. In
Southern Africa specimens are frequently collected in
forests in northern and eastern Transvaal, Swaziland
and Natal. Map 72.
Vouchers: Crosby & Crosby 9140; Magill 3535,
3753, 4892; Vorster 449a.
This species is recognized by its stems appearing
fluted when viewed in cross section, strongly squar-
rose-recurved leaves with serrate margins and the
large crown-shaped papillae of the laminal cells.
Smaller specimens could be confused with L. brachy-
phyllum, especially collections from areas where their
ranges overlap; see note under that species. Both
sporophytes and gemmae have been described for L.
longicaule but neither have been found on Southern
African specimens. Zander (1972) in his revision of
Leptodontium, assigned a specimen ( Wager 1932, US)
to L. longicaule var. microruncinatum (Dus.) Zander.
The variety differs in ‘long-lanceolate leaves, to 6
mm, dentate in the upper \ ; the upper laminal cells
with thickened walls, papillae often obscure or fused
into an irregular lens-shaped cap over each lumen. . .’;
additional specimens have not been seen.
3. Leptodontium brachyphyllum Broth. &
Ther. in Bull. Acad. int. Geogr. bot. 16: 40
(1906); Zander in Bryologist 75: 272 (1972).
Type: Colombia, Bogota, Apollinaire-Marie
s.n., 1904 (FH; H; NY).
Plants medium to large, in loose tufts,
green to dark green, brown to blackish below;
saxicolous. Stems 20-80 mm tall, unbranched,
sparsely tomentose; in section round, central
strand absent, cortical cells large, thickened,
outer cortical cells in 1-3 rows, substereids,
reddish; axillary hairs filamentous, 1 5—16
cells long, hyaline throughout. Leaves crowd-
ed, erect to appressed with weakly twisted
tips dry, recurved wet, not strongly keeled;
ovate-lanceolate, 2,0-2, 5 mm long; apex
Map 72. — • Leptodontium longicaule
x Leptodontium brachyphyllum
Fig. 52.— Leptodontium viticulosoides (1-9): 1. habit, xl; 2. habit, x3; 3. stem in cross section, xl70;
4. leaves, x20; 5. leaf in cross section, x200 ; 6. cells at leaf base (dorsal right side), x 170; 7. upper laminal
cells, x640; 8. leaf apex (dorsal right side, papillae partly shown), x 170; 9. part of capsule mouth with peris-
tome teeth and spores, x 140. L. longicaule (10-16): 10. habit, xl; 11. habit, x2; 12. stem in cross section,
xl70; 13. leaf, x40; 14. leaf in cross section, x200; 15. leaf apex, xl70; 16. upper laminal cells, x640. L.
brachyphyllum (17-24): 17. habit, x 1; 18. habit, x2; 19. stem in cross section, x 170; 20. leaf, x40; 21. leaf
in cross section, x200 ; 22. upper lamina and margin (left side), xl70; 23. upper laminal cells, x640 ; 24.
gemmae, x 170. (1-2, Smook 848; 3-9, Sim 9667; 10-16, Magill 4871; 17-24, Magill 4521).
192
POTTIACEAE
acute; base sheathing; margins recurved to
mid-leaf, serrate above. Costa subpercurrent ;
ventral superficial cells rectangular, smooth,
dorsal superficial cells rectangular, smooth;
in section reniform, guide cells 2-4, large,
ventral stereid band 1-3 cells thick, exposed,
dorsal stereid band 2-4 cells thick, dorsal
surface cells undifferentiated. Upper laminal
cells quadrate to hexagonal or rounded,
moderately thickened, not strongly so in
corners, papillae numerous, appearing scat-
tered over lumen, low, in section forming a
short crown-shaped group over each cell;
basal cells moderately defined, short-rectangu-
lar, thin-walled, papillae in upper base, weak,
scattered. Gemmae on stem, obovoid, to
50 pm long, multicellular, brownish.
Sporophytes not known. Fig. 52: 17-24.
Leptodontium brachyphyllum occurs in Central
and western South America and Southern Africa.
Recent collections have been identified from the
southwestern Cape, eastern Lesotho, Natal, Swazi-
land and eastern Transvaal. Map 72.
Vouchers: Crosby & Crosby 7770; Killick 4214;
Magill 4521, 5788; Vorster 1745.
There remains some doubt about the identity of
these specimens because of a resemblance to L.
longicaule. The short leaves, sparsely tomentose stems,
the slightly bulging, thin-walled, weakly papillose
laminal cells and distinct basal cells, however, clearly
separate specimens of L. brachyphyllum.
Tribe POTTIEAE
Plants very small to large, gregarious to loosely caespitose; terricolous, saxicolous or
occasionally corticolous. Stems erect; central strand present or absent. Leaves broad, orbicular
to elliptical or lingulate to spathulate ; margins plane to revolute, rarely erect, generally entire.
Costa subpercurrent to frequently long-excurrent; in section without ventral stereid band,
occasionally with ventral outgrowths, dorsal stereid band generally strong. Laminal cells
large, smooth, mammillose or papillose, papillae mostly C-shaped.
Capsule cleistocarpic or stegocarpic; peristome present, rudimentary or absent, teeth
mostly long-filamentous, twisted, frequently from high basal membrane; operculum conic,
rostrate or obliquely rostrate.
Key to Genera of Tribe Pottieae
1 Ventral costal surface bearing filaments or lamellae:
2 Costa bearing filaments:
3 Leaf margins broadly involute; costa in section flattened, undifferentiated
1 . Aloina
3 Leaf margins erect or revolute ; costa in section round, with guide cells and stereid
band 2. Crossidium
2 Costa bearing lamellae:
4 Leaves lingulate to broadly lingulate; capsule stegocarpic 3. Pterygoneurum
4 Leaves oval to obovate; capsule cleistocarpic 4. Acaulon
1 Ventral costal surface without filaments or lamellae :
5 Plants small to minute, ephemeral; capsules cleistocarpic:
6 Plants bulbiform; capsules immersed 4. Acaulon
6 Plants mostly taller; capsules emergent or exserted:
7 Capsule just emergent 5. Phascum
7 Capsule long-exserted 6. Pottia
POTTIACEAE
193
5 Plants small to large; capsule stegocarpic:
8 Costa excurrent :
9 Plants very small, bulbiform; leaf margins plane 6. Pottia
9 Plants large, stems generally elongate; leaf margins revolute .8. Tortula
8 Costa mucronate, percurrent or ending below apex:
10 Ventral costa strongly convex; in section ventral surface cells enlarged....
7. Desmatodon
10 Ventral costa not strongly convex:
11 Leaf margins crenulate to serrulate 5. Phascum
11 Leaf margins entire or papillose:
12 Laminal cells mammillose ventrally, flat and smooth to weakly papillose
dorsally; leaf margins plane to involute; plants generally glossy; leaves
involute dry 10. Weisiopsis
12 Laminal cells papillose dorsally and ventrally or rarely smooth; leaf margins
plane or revolute at least below:
13 Plants large; leaves Ungulate to spathulate, 2-3 mm long 8. Tortula
13 Plants small; leaves oval, ovate, oblong or Ungulate, less than 2 mm long:
14 Costa ending below apex 9. Husnotiella
14 Costa percurrent to excurrent 6. Pottia
1. ALOINA
Aloina (C. Miill.) Kindb. in Bih. K. Svenska Vetensk. Akad. Handl. 6: 22 (1882), nom. cons.;
Delgadillo in Bryologist 78: 250 (1975); Broth, in Natiirl. PflFam. 10: 294 (1924); Sim, Bryo.
S. Afr. 233 (1926). Type species: A. aloides (Schultz) Kindb.
Plants small, gregarious, olive-green; terricolous. Stems very short; central strand absent.
Leaves Ungulate, piliferous; margins broadly involute; ventral leaf surface covered with
numerous chlorophyllous filaments. Costa excurrent as a long, smooth, hyaline awn; in
section flattened, undifferentiated. Laminal cells transversely rectangular, smooth.
Dioicous. Capsules long-exserted, cylindrical; peristome teeth linear, papillose, twisted
above short basal membrane; operculum conical; calyptra cucullate, smooth; spores small,
round.
The 9 species of Aloina are mostly restricted to the Northern Hemisphere, although 2 species are South
American endemics. The Southern African species is widely distributed and presently also known from the
United States, Mexico, the Mediterranean Region, Australia and New Zealand.
Aloina bifrons ( De Not.) Delgadillo in
Bryologist 78: 251 (1975). Type: Italy.
Tortula bifrons De Not. in Mem. R. Sci. Torino 40:
305 (1838).
Barbula dregeana C. Miill in Gen. Muse. Fr. 432
(1900). Type: Cape, Namaqualand, Zilverfontein,
Drege s.n., 1830.
Aloina rigida sensu Sim, Bryol. S. Afr. 233 (1926),
non (Hedw.) Limpr. (1888).
Plants small to very small, gregarious,
dark green to olive-green; terricolous. Stems
short, to 3 mm high, simple ; in section without
central strand, cortical cells thin-walled,
gradually smaller toward margins. Leaves
incurved dry, widespreading wet; Ungulate to
ovate-lingulate, 0, 8-2,0 mm long, frequently
constricted above base; apex piliferous; base
sheathing; margins entire to serrulate, broad-
ly involute above base. Costa long-excurrent,
to 1 mm long, fragile, hyaline, smooth;
ventral surface bearing numerous cylindrical
filaments, dorsal superficial cells rounded,
quadrate to rectangular; in section flattened,
undifferentiated, 2-4 cells thick, ventral cells
194
POTTIACEAE
Map 73. — • Aloina bifrons
x Pterygoneurum macleanum
thin-walled, ventral filaments 4-8 cells high,
lower filament cells round, becoming cylind-
rical above, terminal cells elliptical-acute,
strongly thickened apically, dorsal surface
cells strongly thickened. Upper laminal cells
transversely elongate, incrassate, strongly
incrassate dorsally, smooth; basal cells
quadrate to rectangular.
Dioicous. Perichaetia terminal, leaves
clasping, variable in shape. Seta 7-8 mm long,
yellowish red; capsule cylindrical to ovoid-
cylindrical, 1,5-2, 5 mm long, dark red;
peristome spirally twisted, teeth linear above
short basal membrane, yellowish, finely
papillose; operculum conical, erect, to 1 mm
long; spores round, 10-12 pm, finely papillose
to almost smooth. Fig. 53: 1-9.
The widely distributed species, A. bifrons, is
restricted in Southern Africa, to arid and semi-arid
shrublands of the western and southern Cape Province.
Map 73.
Vouchers: Lavranos 15206a; Magill 6075;
Magill & Schelpe 3878a, 3918.
The lingulate leaves with long-excurrent costa,
broadly involute leaf margins and numerous ventral,
photosynthetic filaments characterize this species.
Fig. 53. — Aloina bifrons: 1. habit, x 1; 2. habit,
x5; 3. leaves, x40; 4. leaf in distal cross section,
x 1 70 ; 5. cells at leaf base (left side), x 80 ; 6. marginal
cells and filaments (ventral view), x80; 7. leaf apex,
x80; 8. capsule and detached operculum, x 10; 9.
part of capsule mouth with peristome teeth, xl70.
(1-9, Magill & Schelpe 3896).
POTTIACEAE
195
2. CROSSIDIUM
Crossidium Jur., Laubmfl. Oest. Ungarn 127 (1882), nom. cons.; Delgadillo in Bryologist 78:
269 (1975). Type species: C. squamiferum (Viv.) Jur.
Plants small to minute, scattered or loosely caespitose; terricolous. Stems simple; central
strand weak. Leaves suborbicular to oblong-lanceolate; apex apiculate, subcucullate; margins
erect or revolute, entire or notched. Costa percurrent to mucronate; ventral surface bearing
numerous filaments, dorsal stereid band strong. Upper laminal cells quadrate, weakly thickened.
Sporophyte terminal; capsule cylindrical, reddish; peristome teeth filamentous above
exserted basal membrane; operculum conic to rostrate; calyptra cucullate; spores round,
weakly papillose to smooth.
The genus Crossidium consists of 8 species found primarily in arid or semi-arid regions of both hemi
spheres. In Southern Africa, the genus is found in the northern and western Cape and South West Africa
Namibia. Specimens are rarely collected on crusts of soil under shrubs or in association with rock outcrops.
1 Plants bulbiform; leaves appressed wet or dry; apical cell of costal filaments conical, smooth
1. C. apiculatum
1 Plants larger; leaves spirally twisted around stem dry, erect-spreading wet; apical cell of costal filaments
subglobose, papillose 2. C. spiralifolium
1. Crossidium apiculatum Magill, sp.
nov., speciebus omnibus Crossidii Jur. foliis
suborbicular ibus ad ovalibus, apicibus apicula-
tis, cellulis laminae laevibus minute crassatis,
cellulis apicalibus filamentorum costae laevibus
conicis differt.
Type: Cape, Knersvlakte, near Kobee,
6 km NE of Vanrhynsdorp, along road to
Niewoudtville, Magill & Schelpe 3877 (PRE,
holo. ; BOL; H; MEXU; MO; NY).
Plants minute, bulbiform, scattered,
yellow-green; terricolous. Stems to 1 mm tall,
unbranched; in section round, central strand
very small, weak, cortical cells lax. Leaves
crowded, appressed dry or wet, concave;
suborbicular to oval, 0,6-0, 8 mm long; apex
apiculate, subcucullate; base scarcely differ-
entiated; margins erect, entire or irregularly
notched above. Costa percurrent to mucro-
nate; ventral surface bearing numerous
filaments, dorsal superficial cells rectangular,
smooth; in section oval, guide cells 4, ventral
filaments to 8 cells high, fused at base, apical
cells of filaments conical, smooth, dorsal
stereid band 4 cells thick, dorsal surface cells
undifferentiated. Upper laminal cells rounded,
quadrate to short-rectangular, weakly thick-
ened, smooth; basal cells weakly differen-
tiated, rectangular, hyaline, thin-walled.
Dioicous or cladautoicous. Perichaetia
terminal, leaves to 1 mm long. Seta 2, 5-3, 5
mm long, yellow; capsule elliptical, 1 mm
Map 74. — • Crossidium spiralifolium
x Crossidium apiculatum
long, dark red to yellowish red; peristome
with basal membrane projecting above
capsule mouth, to 50 pm high, teeth short,
irregularly cleft, 200-300 pm high, straight,
strongly papillose; operculum conic to
rostrate, 0,2-0, 5 mm long, cells weakly
twisted counter-clockwise; spores subround,
16-23 pm, essentially smooth. Fig. 54:
1-10.
Endemic to Southern Africa, C. apiculatum is
found on crusts of sand under low shrubs in Namaqua-
land in the northwestern Cape. Map 74.
Vouchers: Magill & Schelpe 3881, 3890, 3891,
3892.
196
POTTIACEAE
Fig. 54. — Crossidium apiculatum (1-10): 1.
habit, x 1 ; 2. habit, showing plants with conical and
rostrate opercula, x 10; 3. stem in cross section,
xl70; 4. leaves, x40; 5. leaf in distal cross section,
x 170; 6. basal leaf cells, x 170; 7. upper laminal cells,
x 640; 8. leaf apex (dorsal surface), x 170; 9. capsule,
x 50; 10. part of capsule mouth with peristome teeth,
x230. C. spiralifolium (11-17): 11. habit, xl; 12.
habit, xlO; 13. leaf, x30; 14. leaf in distal cross
section, x 170; 15. upper laminal cells, x640; 16.
leaf apex (dorsal surface), xl70; 17. part of capsule
mouth with peristome teeth, x 100. (1—10, Magill &
Schelpe 3877 ; 1 1-17, Smook & Harding 705).
POTTIACEAE
197
Crossidium apiculatum differs from the other
species of Crossidium by its apiculate leaf apices,
smooth, conical apical cells of the costal filaments and
suborbicular, appressed leaves.
2. Crossidium spiralifolium Magill, sp.
nov., C. rosei Williams foliis oblongo-lanceo-
latis mucronatis, cellulis apicalibus filamen-
torum costae subglobosis et papillosis simile,
sed cellulis laminae laevibus, peristomio bene
evoluto et sporibus minoribus dijfert.
Type: Cape, Prieskapoort, c. 14 km
from Prieska on road to Vosburg, on soil,
Smook & Harding 705 (PRE, holo. ; H;
MEXU; MO; NY).
Plants small, loosely caespitose, dark
green above, brownish below; terricolous.
Stems 2 mm tall, unbranched; in section with
central strand, cortical cells thin-walled, in
4-5 rows, outer row smaller, reddish. Leaves
appressed, spirally twisted around stem dry,
erect-spreading wet; oblong-lanceolate, 1,2
mm long; apex acute; base scarcely differen-
tiated; margins narrowly revolute, entire.
Costa mucronate; ventral surface bearing
numerous filaments, dorsal superficial cells
rectangular, smooth; in section subround,
guide cells 6, ventral filaments 4-6 cells
high, apical cells of filaments subglobose,
with 2-4 papillae, filaments frequently
branched above, dorsal stereid band strong,
to 8 cells thick, dorsal surface cells undif-
ferentiated. Upper laminal cells quadrate to
angular, thin-walled, smooth; in section
dorsal surface strongly thickened; basal cells
weakly differentiated, quadrate to short-
rectangular, hyaline, thin-walled.
Dioicous or cladautoicous. Perichaetia
terminal, leaves to 1,5 mm long. Seta to
10 mm long, yellow; capsule cylindrical,
1,2-1, 5 mm long, reddish yellow; peristome
with basal membrane projecting above
mouth, to 50 pm high, teeth irregularly cleft,
long-filiform, 500 //m high, twisted counter-
clockwise 1-2 times, yellowish red, papillose;
operculum long-conic, to 0,6 mm long, cells
twisted counter-clockwise; spores round to
subround, 18-20 pm, weakly papillose.
Fig. 54: 11-17.
Endemic to Southern Africa, this species is
presently known from the shrublands of the central
Cape Province and southern South West Africa/
Namibia. Map 74.
Voucher: Hardy 4864a.
This species differs from C. apiculatum in size,
leaf shape and costal filament morphology. Crossi-
dium spiralifolium is separated from other species of
the genus by the combination of mucronate leaves,
smooth leaf cells, the terminal cell of costal filaments
being subglobose and papillose, and the well developed
peristome.
3. PTERYGONEURUM
Pterygoneurum Jur., Laubmfl. Oest. Ungarn 95 (1882), nom. cons.; Broth, in Natiirl. PflFam.
10: 292 (1924); Sim, Bryo. S. Afr. 220 (1926); Wareham in Grout, Moss FI. N. Amer. 1: 208
(1939). Type species: P. cavifolium Jur.
Plants small, forming dense cushions, dark green; saxicolous. Stems simple; central
strand absent. Leaves lingulate; apex obtuse, subcucullate; margins broadly involute. Costa
short-excurrent; ventral surface bearing high, plate-like, chlorophyllous lamellae. Laminal
cells quadrate, smooth.
Autoicous. Perichaetia terminal; capsule exserted, subglobose; peristome rudimentary or
absent; operculum short-rostrate; calyptra cucullate, large; spores large, papillose.
The 8 species of Pterygoneurum are mostly restricted in distribution. Three species occur sporadically in the
Northern Hemisphere and one of these also reaches Australia. Four other species are known from Africa, 3
from the northern part of the continent and 1 is endemic to Southern Africa.
Pterygoneurum macleanum Warnst. in Plants small, in dense cushions, olive-
Hedwigia 58: 69 (1916); Broth, in Natiirl. green, brownish below; saxicolous. Stems to
PflFam. 10: 292 (1924); Sim, Bryo. S. Afr. 220 5 mm high, simple; in section round, central
(1926). Type: Cape, Graaff-Reinet, MacLea strand absent, but central cells often coloured,
sub Rehmann 461 (PRE!). cortical cells lax, somewhat thickened at
198
POTTIACEAE
Fig. 55.— Pterygoneurum macleanum: 1. habit,
xl; 2. habit, xlO; 3. stem in cross section, xlOO;
4-5. leaves, x40; 6. leaf in distal cross section, x240;
7. cells at leaf base, x 170; 8. leaf apex, x 170. (1-8,
Rehmann 461).
margins. Leaves incurved-appessed or slightly
contorted dry, erect wet; lingulate to ovate-
lingulate, 1,0-1, 5 mm long; apex obtuse;
margins broadly involute above. Costa short-
excurrent to mucronate; ventral surface with
4 high, undulate lamellae, dorsal superficial
cells long-rectangular, strongly incrassate; in
section subround, guide cells 4, each bearing a
plate-like lamella, to 16 cells high, dorsal
stereid band to 6 cells thick, dorsal surface
cells substereids. Upper laminal cells quadrate,
incrassate, smooth; basal cells similar or
slightly larger, thin-walled; lamellae cells
quadrate, thin-walled, chlorophyllous.
Autoicous. Perichaetial leaves undifferen-
tiated. Seta 1 ,0-1,2 mm long, yellowish; cap-
sule subglobose, to 1 mm long, gymnosto-
mous; operculum rostrate, to 0,7 mm long;
calyptra cucullate, 2 mm long, covering cap-
sule; spores large, 35-38 pm, papillose. Fig.
55: 1-8.
Endemic to Southern Africa, the species is known
from the semi-arid shrublands of the central Cape
Province. Map 73.
Voucher: Type only.
Pterygoneurum macleanum bears some resem-
blance to species of Crossidium but can be distin-
guished from them by the plate-like lamellae and
gymnostomous capsule. Ventral costal lamellae are
also known in Acaulon and several genera of the
Polytrichaceae.
4. ACAULON
Acaulon C. Mull, in Bot. Ztg 5: 99 (1847); Broth, in Natiirl. PflFam. 10: 283 (1924); Scott &
Stone, Moss. S. Aust. 176 (1976). Type species: A. muticum (Hedw.) C. Mull.
Plants very small, bulbiform, hyaline or reddish green, gregarious; growing on sand or
clay. Stems simple; central strand absent. Leaves imbricate wet or dry, concave; broadly oval
to obovate; apex acute, frequently hyaline; margins plane or revolute, entire or dentate at
apex. Costa aristate, hyaline or reddish; in section round, guide cells 2, ventral cells present
or absent, sometimes producing plate-like lamellae, dorsal cells stereids or substereids. Laminal
cells quadrate, rhomboidal or hexagonal, smooth or infrequently weakly papillose; basal
cells rectangular.
Paroicous or autoicous. Perichaetial leaves ± larger, similar to upper leaves. Seta very
short; capsule cleistocarpic, immersed, globose-apiculate; exothecial cells of various shapes,
irregularly arranged; calyptra minute, mitrate; spores round, granulate to minutely spinose.
The 18 species of Acaulon are infrequently collected in temperate regions of both hemispheres. In Southern
Africa the genus is known from the arid and semi-arid regions of the western Cape and South West Africa/
Namibia. The genus is closely related to Phascum and differences between the two are discussed there.
POTTIACEAE
199
1 Plants green to reddish green; leaf awn reddish yellow 3. A. rufochaete
1 Plants hyaline above, light green below; leaf awns hyaline:
2 Upper ventral costal surface with short plate-like lamellae, 2-6
cells high; leaf margins plane 1. A. leucochaete
2 Upper ventral costal cells enlarged, swollen, not forming lamellae;
leaf margins revolute 2. A. recurvatum
1 . Acaulon leucochaete Stone in J. Bryol.
9: 217 (1976). Type: Australia, Victoria,
Boundary Bend, Stone 1548b (MEL, holo.;
MELU; BM).
Plants small, hyaline above, light green
below; terricolous. Stems 0,2-0, 3 mm tall; in
section round, central strand absent or with
1- 3 smaller cells, cortical cells lax, reddish
toward margin. Leaves imbricate and with
flexuose awns wet or dry, concave; broadly
oval to oblong, 0 , 5-1 , 0 mm long ; apex acute,
hyaline; margins plane, entire below, irregu-
larly dentate near apex. Costa long-excurrent
as a smooth, hyaline awn, to 1 mm long in
upper leaves; upper ventral surface with
lamellae, dorsal superficial cells rectangular,
smooth; in section round, guide cells 2, ven-
tral, each bearing a short, plate-like lamella,
2- 6 cells high, dorsal stereid band 2 cells thick,
cells frequently substereids, dorsal surface
cells substereids, outer walls strongly
thickened. Upper laminal cells rhomboidal to
hexagonal, thin-walled, smooth; marginal
cells narrower, forming weak border in upper
leaf ; basal cells rectangular.
Paroicous. Perichaetial leaves not diffe-
rentiated. Seta very short, 0,2-0, 3 mm long;
capsule cleistocarpic, globose with minute
apiculus, to 1 mm long, reddish yellow; exo-
thecial cells quadrate to short-rectangular;
calyptra small, mitrate; spores round, 23-25
pm, strongly incrassate, granulate to almost
smooth, frequently with 3-4 large oil bodies.
Fig. 56: 1-11.
Acaulon leucochaete is known from Australia and
Southern Africa. Growing under low shrubs on sandy
or clay soils, specimens have recently been collected
in the succulent shrublands, between Clanwillian and
Springbok, in the northwestern Cape. Map 75.
Vouchers: Magill & Schelpe 3883, 3895, 3914;
Schelpe 7766.
This species was recently described from Boun-
dary Bend in Victoria, Australia, by Stone (1976)
along with a closely related species, A. chrysacanthum
Stone. The presence of A. leucochaete in Southern
Africa is a clear indication of the close relationship of
the two bryofloras. Acaulon leucochaete is separated
from the other species by ventral costal lamellae and
plane leaf margins.
2. Acaulon recurvatum Magill, sp. nov.,
habitu et colore A. leucochaete Stone simile,
sed marginibus foliorum integris, reflexis ad
revolutis, cellulis laminae infirme papillosis, et
cellulis ventralibus superficialibus costae ampli-
ficatis papillosis differt.
Type: Cape, Namaqualand, Springbok,
on soil 10 km E of Springbok, on Pofadder
Road, Schelpe 77 17 (BOL, holo.; MO; PRE).
Plants small, hyaline above, yellow-green
below; terricolous. Stems 0, 1-0,2 mm tall; in
section without central strand, cortical cells
lax, marginal cells reddish brown. Leaves
imbricate with contorted awns wet or dry,
concave; broadly oval to obovate, 1,0- 1,5
mm long; apex acute, frequently hyaline;
margins revolute above base, entire to denti-
culate. Costa excurrent as long, smooth,
hyaline awn, 0,5-0, 8 mm in upper leaves;
ventral superficial cells lax, bulging, quadrate
to short-rectangular, with 1-2 weak papillae,
dorsal superficial cells elongate, smooth; in
section round, guide cells indistinct, ventral
surface cells enlarged, swollen, 4-6 cells in
single row, papillose, dorsal stereid band 5-6
cells thick, some cells substereids, dorsa
Map 75. — • Acaulon recurvatum
x Acaulon leucochaete
A Acaulon rufochaete
200
POTTIACEAE
POTTIACEAE
201
surface cells undifferentiated. Upper laminal
cells quadrate, incrassate, smooth; median
leaf cells quadrate to hexagonal, finely papil-
lose, with 1-3 C- or O-shaped papillae over
dorsal lumen; basal cells larger, short-
rectangular.
Paroicous. Perichaetial leaves broader;
margins reflexed to revolute; apical cells
smooth, rhomboidal, hyaline, thin-walled.
Seta to 0,3 mm long; capsule cleistocarpic,
globose-apiculate, 0,9-1 ,0 mm long, reddish
yellow; exothecial cells short-rectangular to
quadrate; calyptra minute, mitrate, 0,4 mm
long; spores round, 22-25 pm, granulate. Fig.
56: 12-18.
Endemic to Southern Africa, this species occurs
in the succulent Karoo of Namaqualand, in the north-
western Cape and fynbos of the southern and south-
western Cape. Map 75.
Voucher: Magill & Schelpe 4046; Muir 3779a.
Similar to A. leucochaete, but differs in the en-
larged, papillose, ventral superficial cells of the costa
and the absence of lamellae. The upper leaf margins
also differ in being entire and reflexed to revolute.
3. Acaulon rufochaete Magill, sp. nov.,
habitu et colore A. chrysacantho Stone simile,
sed foliis marginibus recurvatis, costa sine
lamellis ventralibus, cellulis laminae interdum
subtiliter papillosis et sporis majoribus differt.
Type: Cape, Knersvlakte, on soil 6 km
NE of Vanrhynsdorp, near turnoff to Kobee,
Magill & Schelpe 3877c (PRE, holo.; BOL;
H; MO; NY).
Plants very small, scattered, green to
reddish green ; terricolous. Stems 0, 1-0,2 mm
tall; in section without central strand, cortical
cells lax. Leaves imbricate dry, imbricate with
erect apices and reflexed awns wet; broadly
oval to oblong, 0 , 8-1 , 5 mm long ; apex acute,
chlorophyllous; margins revolute above,
entire. Costa excurrent as smooth, reddish
yellow awn, 0, 1-0,5 mm long, ventral and
dorsal superficial cells rectangular, smooth; in
section round, guide cells exposed ventrally,
2-4, weakly thickened, dorsally cells incras-
sate, 2-4 cells thick. Upper laminal cells
quadrate, rhomboidal or short-rectangular,
thin-walled or weakly thickened, smooth or
rarely with 1-2 minute, blunt papillae over
lumen; basal cells short-rectangular, thin-
walled, smooth.
Autoicous. Perichaetial leaves oblong to
oval, acute, concave ; costa excurrent, reddish ;
laminal cells short-rectangular to rhombic.
Seta 0, 1-0,2 mm long; capsule cleistocarpic,
globose-apiculate, 0,5 mm long, reddish
yellow, darker above; exothecial cells triangu-
lar, rectangular or angular-elongate; calyptra
minute, mitrate, 0,3 mm long; spores round,
37-42 pm, minutely spinose. Fig. 56: 19-25.
Presently known only from the western Cape,
the species is infrequently collected on sandy soil
under low shrubs between Clanwilliam and Vanrhyns
Pass. Map 75.
Vouchers: Magill & Schelpe 3878, 3897, 3912.
This species is recognized by its long reddish awn,
more or less smooth leaf cells and large, minutely
spinose spores.
Insufficiently Known Species
Acaulon capense C. Mull, in Bot. Ztg 14: 415
(1856). Sphaerangium capense (C. Mull.) Jaeg. in Verh.
St Gall, naturw. Ges. 1871-72: 336 (1873). Type:
Cape, Swellendam, Pappe s.n. The type has not been
seen. Sim (1926) referred this species to A. muticum
(Hedw.) C. Mull, without seeing Pappe’s specimen.
Acaulon sphaericum Shaw in Cape Monthly Mag.
17: 313 (1878). Type: Cape, Graaff-Reinet, McLea s.n.
Sim (1926) refers this species to A. muticum (Hedw.)
C. Mull, without having seen the type; however, the
description could also apply to an Ephemerum species.
The Sim correspondence (PRE) indicates that the
Shaw collection was destroyed sometime after his
death. Attempts to locate the specimens in Southern
Africa have been unsuccessful and duplicates were not
located in the European herbaria likely to house
Shaw specimens.
Fig. 56. — Acaulon leucochaete (1-11): 1. habit, xl; 2. habit, x20; 3. stem in cross section, X100; 4.
lower stem leaf, x40; 5. upper leaves, x40; 6. leaf in distal cross section, xl70; 7. basal leaf cells, x!70;
8. cells at leaf apex (dorsal surface), xl70; 9. capsule with calyptra, x40; 10. exothecial cells and stomata
at capsule base with spores, x80; 11. spore, x400. A. recurvatum (12-18): 12. habit, xl; 13. habit, x20;
14. leaves, x40; 15. leaf in distal cross section, xl70; 16. upper laminal cells, x400; 17. cells at leaf apex
(dorsal surface), x 170; 18. spore, x 500. A. rufochaete (19—25) : 19. habit, x 1 ; 20. habit, x20; 21. leaves, x40;
22. leaf in distal cross section, x 170; 23. upper laminal cells, x400; 24. cells at leaf apex (dorsal surface), x 170;
25. perichaetial leaf, x40. (1-11, Magill & Schelpe 3900; 12-18, Schelpe 7717; 19-25, Magill & Schelpe 3912).
202
POTTIACEAE
5. PHASCUM
Phascum Hedw., Spec. Muse. 19 (1801); Broth, in Natttrl. PflFam. 10: 284 (1924); Sim, Bryo.
S. Afr. 218 (1926); Grout, Moss FI. N. Amer. 1 : 195 (1939). Lectotype species: P. cuspidatum
Hedw., vide Grout, Moss FI. N. Amer. 1: 195 (1939).
Plants small, gregarious or in small groups, yellowish green to reddish green; terricolous.
Stems to 5 mm high, generally smaller; central strand absent. Leaves little contorted dry,
erect to erect-spreading wet; obovate, oblong or oval; apices acute; margins plane or revolute,
entire or serrate. Costa short- or long-excurrent, smooth; in section with dorsal stereid or
substereid band only. Laminal cells hexagonal, quadrate or rectangular, smooth or finely
papillose.
Autoicous. Capsule cleistocarpic, emergent or exserted, elliptical, rostrate; calyptra
cucullate; spores round to oval, almost smooth or verruculate.
A genus of c. 20 species, Phascum is widespread although uncommon in temperate regions. In Southern
Africa the genus is best represented in the drier areas of the western Cape and southern South West Africa/
Namibia. Phascum leptophyllum is known from more mesic locations in the eastern Cape, eastern Transvaal and
Zimbabwe.
The genus is closely related to Acaulon and the two genera are frequently collected together in Namaqua-
land. The classic diagnostic characters appear to overlap in Southern African species, as is the case in Australia,
(Scott & Stone, 1976). The size of the capsule beak, curvature of leaf margin and papillosity of leaf cells, although
useful characters, will not easily separate Southern African taxa. Phascum is defined here by its narrower,
generally plane-margined, erect or spreading leaves, and its emergent capsule and cucullate calyptra. The
leaves of the Southern African species of Acaulon are broad, concave and enclose the capsule, giving a bulbiform
appearance to the plants. The calyptra is very small and mitrate.
1 Plants 2-5 mm tall; leaf margins irregularly serrate; costa
short-excurrent
1 Plants generally less than 1 mm tall; leaf margins entire; costa
excurrent as a long, reddish or yellowish awn
1. Phascum peraristatum C. Mull, in
Flora, Jena 71: 3 (1888); Broth, in Natiirl.
PflFam. 10: 284(1924). Type: Cape, Somerset
East, Boschberg, MacOwan s.n., 1882.
Plants small, gregarious, reddish green;
terricolous. Stems erect, 0, 5-1,0 mm tall; in
section without central strand, cortical cells
large, weakly thickened, smaller at margin,
yellowish red. Leaves erect dry, erect-
spreading wet; oblong-obovate to oval,
0,8-1, 2 mm long; apex acute, sometimes
abruptly so; margins revolute above base,
entire. Costa excurrent, awn 0, 1-1,0 mm
long, reddish to yellowish; ventral super-
ficial cells quadrate, papillose to near apex,
absent above and exposing elongate, smooth
cells, dorsal superficial cells elongate, smooth;
in section round, guide cells 2, small, incras-
sate, ventral cells in single layer, 2-4, large,
papillose, dorsal stereid band 3 cells thick,
dorsal surface cells frequently substereids.
Upper laminal cells quadrate to subhexagonal,
sometimes short-rectangular, with 2-6 low,
2. P. leptophyllum
1. P. peraristatum
C-shaped papillae over lumen, rarely smooth;
basal cells larger, quadrate to rectangular or
some angular, smooth.
Map 76. — • Phascum peraristatum
X Phascum leptophyllum
POTTIACEAE
203
Fig. 57. — Phascum peraristatum (1-11): 1.
habit, x 1 ; 2. habit, xlO; 3. stem in cross section,
xl70; 4. lower leaf, x40; 5. upper leaves, x40; 6.
leaf in cross section, X 170; 7. cells at leaf base (right
side), xl70; 8. upper laminal cells, x640; 9. cells at
leaf apex, x 170; 10. capsule with attached perichaetial
leaf, x40; 11. calyptra, x40. P. leptophyllum (12-17):
12. habit, xl; 13. upper stem, xlO; 14. leaf x40;
15. leaf in cross section, xl70; 16. upper laminal
cells, x640; 17. leaf apex, xl70. (1-11, Magill &
Schelpe 4011a; 12-17, Vorster 2198).
3
204
POTTIACEAE
Autoicous, frequently polysetaceous.
Perichaetial leaves undifferentiated. Seta very
short, 0,2-0, 3 mm long; capsule cleistocar-
pic, emergent, reddish yellow, short-elliptical
to subglobose, rostrate, 0,4-0, 5 mm long,
exothecial cells quadrate at apex and base,
median cells narrowly oblong to triangular;
calyptra cucullate, small, 0,4 mm long; spores
round to oval, 25-33 /an, smooth to irregu-
larly granulate. Fig. 57: 1-11.
Originally described from the eastern Cape
Province, P. peraristatum has recently been found at
several locations in the dry shrublands of the western
Cape. Map 76.
Vouchers: Magill & Schelpe 4011a; Schelpe 7765,
7781.
This species may be confused, vegetatively, with
Acaulon. Sporophytically, P. peraristatum is distinct in
the exposed, elliptical-rostrate capsules, differentiated
exothecial cells and cucullate calyptra.
The species was described as having smooth leaf
cells, however all subsequent collections examined
from Southern Africa, have several fine, C-shaped
papillae on the upper laminal cells. Since these
papillae were detected on specimens mounted in
Hoyer’s medium, it was believed that the papillae
had been overlooked when the species was described.
Unfortunately the type has not been located.
A specimen recently examined from Kenya
(Comm. Townsend, 1981) is similar to the Southern
African specimens except that it has smooth leaf
cells. It may, therefore be more similar to MacOwan’s
collection than the papillose specimens from the
western Cape. The taxon is considered to include
both “facies”, however the papillose specimens may
represent an undescribed variety.
2. Phascum leptophyllum C. Mull, in
Flora, Jena 71: 6 (1888); Broth, in Natiirl.
PflFam. 10: 284 (1924); Sim, Bryo. S. Afr. 218
(1926). Type: Cape, Somerset East, Bosch-
berg, MacOwan s.n., 1882.
Plants small, gregarious, light green;
terricolous. Stems 2-5 mm high; in section
round, central strand absent, cortical cells lax,
outer row smaller. Leaves somewhat distant,
weakly contorted dry, erect-spreading wet;
narrowly elliptical to obovate, 1, 5-3,0 mm
long; apex acute; margins plane, irregularly
crenulate to serrate above base. Costa short-
excurrent; ventral and dorsal superficial
cells short-rectangular, smooth; in section
round, guide cells 2, small, thickened, ventral
surface cells 2, thin-walled, bulging, dorsal
cells weakly incrassate, 3 cells thick. Upper
laminal cells hexagonal to subhexagonal,
quadrate or rectangular, smaller toward
margin, smooth; basal cells quadrate to
short-rectangular.
Sporophyte not seen, but described by
Muller as: Seta short; capsule emergent,
turgid-elliptical with distinct, oblique beak,
brownish. Fig. 57: 12-17.
Phascum leptophyllum was described from Somer-
set East in the shrublands of the central Cape Pro-
vince. Sim made additional collections from Victoria
Falls, Zimbabwe, and more recently specimens have
been collected in Botswana, Natal and the south-
western Cape. Map 76.
Vouchers: Sim 8771, 8857; Magill 5062; Van
Zanten 7608116.
The plants are very large for Phascum and may
be more properly placed in Pottia. Additional sporo-
phytes must be found in order to assess proper
generic position.
6 POTTIA
Pottia ( Reichenb .) Fuernr. in Flora, Jena 12 (2 Erg.): 10 (1829); Broth, in Natiirl. PflFam. 10:
289 (1924); Wareham in Grout, Moss FI. N. Amer. 1: 197 (1939); Sim, p.p., Bryo. S. Afr.
219 (1926). Lectotype species: P. truncata (Hedw.) B.S.G., vide Wareham ex Grout, Moss
FI. N. Amer. 1: 197 (1939).
Plants small to minute, gregarious or in loose cushions, yellowish, reddish or dark green;
terricolous. Acaulescent or stems to 2 mm tall; central strand absent or not well defined,
Fig. 58. — Pottia namaquensis (1-11): 1. habit, xl; 2. habit, X20; 3. leaves, x40; 4. leaf in cross section,
x230; 5. basal leaf cells, x 170; 6. upper laminal cells, x640; 7. cells at leaf apex, X 170; 8. perichaetial leaf,
x40; 9. capsule, x30; 10. calyptra, x30; 11. part of capsule mouth with peristome teeth and spores, x 175.
P. macowaniana (12-20): 12. habit, xl; 13. habit, x20; 14. leaves, x40; 15. leaf in cross section, x230; 16.
basal leaf cells (dorsal surface), Xl70; 17. upper laminal cells, x370; 18. leaf apex (left dorsal side), xl70;
19. capsule, x20; 20. part of capsule mouth with peristome teeth, x250. P. subplanomarginata (21-29): 21.
habit, x 1 ; 22. habit, x 10; 23. stem in cross section, x230; 24. leaves, X40; 25. leaf in cross section, x230;
26. basal leaf cells, xl70; 27. leaf apex, Xl70; 28. upper laminal cells, x640; 29. perichaetial leaf, x40.
(1-1 1, Oliver 7208; 12-20, MagiinSSSc; 21-29, Wager 1570c).
POTTIACEAE
205
fogjiTfia
206
PoTTIACEAE
cortical cells lax. Leaves erect to spreading wet; oval, ovate or oblong; apex acute to obtuse;
margins plane, erect or revolute. Costa subpercurrent to long-excurrent; in section guide cells
small, ventral surface cells similar to laminal cells, dorsal stereid band small. Laminal cells
quadrate, hexagonal or rectangular, finely papillose; basal cells generally larger, quadrate to
rectangular.
Autoicous. Perichaetia terminal, leaves undifferentiated. Capsule stegocarpic or cleisto-
carpic, emergent or exserted, oval to ovate or short-cylindrical, peristome absent or, when
present, rudimentary, consisting of short, cleft or perforated teeth, frequently only lower base
of teeth present, finely papillose, yellowish; operculum conic; calyptra cucullate; spores
round to angular, smooth, granulate or verruculate.
The genus is widely distributed in temperate regions, although the c. 60 species tend to be restricted in their
individual distributions. In Southern Africa, Pottia is known only from the Cape and southern South West
Africa/Namibia. The four species are rarely collected, and occur on sandy soil in association with other epheme-
ral species of Acaulon, Phascum, Aloina and Crossidium.
I Costa long-excurrent; peristome developed 1. P. namaquensis
1 Costa short-excurrent or percurrent; peristome absent or rudimentary:
2 Capsule cleistocarpic; costa percurrent; leaves oblong to Ungulate 4. P. splachnoides
2 Capsule stegocarpic, peristome rudimentary; costa short-excurrent:
3 Leaves oblong-lanceolate to oblong-elliptical; plants in loose groups; spores granulate, 25-32 pm
3. P. subplanomarginata
3 Leaves broadly ovate to oval; plants gregarious; spores verruculate, 17-22 pm. ... 2. P. macowaniarta
1. Pottia namaquensis Magill, sp. nov., in
genera bene distinta speciebus ceteris Pottiae
( Reichenb .) Fuernr., capsula emergente, parva
cylindracea, calyptra scabrosa, seta minima et
foliis late ovalibus costis longe excurrentibus
dignoscenda.
Type: Cape, Namaqualand, Messelpad
Road between Taaiboskraal and Nuweputs,
south slope, damp, open ground, Oliver 7208
(PRE, holo.; MO).
Plants very small, acaulescent or nearly
so, 1 -2 mm high, bulbiform, dark green, grega-
rious; terricolous. Leaves appressed dry,
erect-spreading wet, concave; broadly oval,
0,5-1 ,0 mm long; apex rounded; margins
plane, entire. Costa long-excurrent as a
smooth, hyaline awn, to 1 mm long; ventral
superficial cells quadrate, papillose, dorsal
superficial cells long-rectangular, smooth; in
section with 4 large guide cells, ventral cells in
two rows, inner cells smaller than guide cells,
surface cells similar to laminal cells, papillose,
dorsal stereid band 4-5 cells thick, dorsal
surface cells undifferentiated. Upper laminal
cells quadrate to subhexagonal, papillae 1-3
per cell, C- or O-shaped; basal cells quadrate,
larger, smooth, hyaline.
Dioicous. Perichaetial leaves clasping
capsule; elliptical, 2 mm long including awn.
Seta very short, to 0,3 mm long; capsule
emergent, short-cylindrical, 0,6 mm long;
exothecial cells rectangular, cells at mouth
narrow, vertically elongate, in 1-3 rows;
peristome short, 0,15 mm high, somewhat
irregular, teeth triangular, cleft or perforated,
Map 77. — • Pottia subplanomarginata
X Pottia namaquensis
POTTIACEAE
207
above, papillose; operculum rostrate, 0,4 mm
long; calyptra broadly conic, long-rostrate,
0,7 mm long, spinose or with short hairs;
spores round, 17-18 p m, smooth. Fig. 58:
1-11.
Endemic to Southern Africa, P. namaquensis was
collected in the dwarf succulent shrublands of Nama-
qualand and the Richtersveld in the northwestern
Cape. Map 77.
Vouchers: Oliver, Tdlken & Venter 651a, PRE-
CH12888, PRE-CH12897.
The species is recognized by the very short stem,
the broadly oval, aristate leaves and emergent, short-
cylindrical capsule.
2. Pottia macowaniana C. Mull, in Hed-
wigia38: 98(1899); Broth, in Natiirl. PflFam.
1 : 423 (1902); Sim, Bryo. S. Afr. 219 (1926).
Type: Cape, Small Fish River, MacOwan s.n.,
1878.
? Pottia verrucosa Rehm. ex Warnst., p.p., in
Hedwigia 58: 140 (1916). Type: Cape, Graaff-Reinet,
McLea sub Rehmann 460 (B).
Plants minute, gregarious, green ; terrico-
lous. Stems to 0,5 mm tall; in section central
cells very thin-walled, cortical cells lax. Leaves
imbricate dry, patent wet; broadly ovate to
oval, 0,8-1, 8 mm long; apex obtuse to
rounded, mucronate to apiculate; margins
revolute above base, entire. Costa short-
excurrent, smooth; ventral superficial cells
quadrate to short-rectangular, papillose, dor-
sal superficial cells rectangular, smooth; in
section round, guide cells 2, small, ventral
surface cells similar to lamina cells, papillose,
dorsal stereid band 2-4 cells thick, dorsal
surface cells substereids, sometimes lateral
cells larger, papillose. Upper laminal cells
quadrate to hexagonal, some short-rectangu-
lar, papillose, 2-4 C-shaped papillae over
lumen ; basal cells larger, rectangular to
quadrate, smooth.
Autoicous. Antheridia axillary on upper
stem ; perichaetia terminal, leaves undifferen-
tiated. Seta 2, 0-3, 5 mm long, yellowish;
capsule cylindrical to oval, 1 ,0-1 ,2 mm long;
peristome rudimentary, exserted above
mouth, light yellow, faintly papillose; oper-
culum short-conic; calyptra cucullate, to 1-2
mm long; spores irregular, rounded to
triangular, 17-22 pm, weakly verruculate.
Fig. 58: 12-20.
Originally described from the fynbos of the
southeastern Cape Province; recent collections have
been made in the dwarf succulent shrublands of the
western and northern Cape. Map 78.
Vouchers: Barnard PRE-CH5760; Booi 126;
Magill & Schelpe 3879, 3882, 3908.
These very small, bulbous plants are recognized
by the broadly ovate to oval leaves, percurrent costa
and long-exserted capsules. They could be confused
with papillose forms of P. splachnoides when sterile,
but the more differentiated costal anatomy and
recurved margins help to distinguish the species.
3. Pottia subplanomarginata Dix. in
Trans. R. Soc. S. Afr. 18: 253 (1930). Type:
Cape, Stellenbosch, Wager 671 (BM, holo. !;
PRE!).
Plants small, in loose cushions, yellowish
to reddish green; terricolous. Stems 1-2 mm
tall; in section round, central cells very thin-
walled, cortical cells lax, yellowish. Leaves
weakly contorted dry, spreading wet; oblong-
lanceolate to oblong-elliptical, 1,2-1, 5 mm
long; apex acute, cuspidate; margins narrow-
ly revolute to plane, entire. Costa short-
excurrent, awn to 0,1 mm long, smooth;
ventral superficial cells quadrate to short-
rectangular, papillose, dorsal superficial cells
rectangular, smooth; in section round, guide
cells 2, ventral surface cells larger, thickened,
papillose, dorsal stereid band small, 1-3 cells
thick, dorsal surface cells rounded, thickened,
smooth. Upper laminal cells hexagonal to
rectangular, papillose, 4-6 C-shaped papillae
over lumen; marginal cells narrower, quadrate
to rectangular, papillose; basal cells large,
rectangular, smooth.
Map 78.— • Pottia macowaniana
x Pottia splachnoides
208
POTTIACEAE
Autoicous. Seta 2-4 mm long, yellowish;
capsule ovoid to cylindrical, 0,8 mm long;
exothecial cells rectangular, 2 rows at mouth
quadrate; peristome very rudimentary to
absent, consisting of 1-2 lightly papillose
cells at mouth; operculum rostrate, deci-
duous; calyptra cucullate; spores angular,
25-32 //m, granulate. Fig. 58: 21-29.
Known only from Southern Africa, P. subplano-
marginata is found in shrublands of the western and
eastern Cape Province and South West Africa/
Namibia. Map 77.
Vouchers: Magill & Schelpe 3877a, 3884; Wager
7837.
The relatively long, oblong leaves and short-
excurrent costa of P. subplanomarginata will separate
it from the other Southern African species.
The description and illustration of P. verrucosa
strongly suggest that the type (B) was P. macowaniana,
however, the specimen at NH! is P. subplanomargi-
nata.
4. Pottia splachnoides ( Hornsch .) Broth.
in Natiirl. PflFam. 1 : 423 (1902). Type: Cape,
Ranker Bay, Bergius s.n. (G!).
Phascum splachnoides Hornsch. in Hor. Phys.
Berol. 57 (1820).
Plants small, gregarious or in small
groups, yellowish green; terricolous. Stems
1-2 mm high; in section round, central strand
absent, cortical cells lax. Leaves weakly
crisped dry, erect wet; oblong to lingulate,
0,8-1, 8 mm long, weakly concave; apex
acute; margins plane to erect, entire. Costa
percurrent to subpercurrent; ventral superfi-
cial cells quadrate, smooth or papillose, dorsal
superficial cells rectangular, smooth or papil-
lose; in section round, guide cells small, gene-
rally undifferentiated, central cells small, in-
crassate, ventral and dorsal surface cells
larger, thin-walled. Laminal cells turgid, sub-
quadrate to short-rectangular, thin-walled,
almost smooth or with 4-8 fine, C-shaped
Fig. 59. — Pottia splachnoides: 1. habit, xl; 2.
habit, x20; 3. leaves, x50; 4. leaf in cross section
(smooth-celled form), x435; 5. leaf in cross section
(papillose-celled form), x435; 6. basal leaf cells,
xl70; 7. upper laminal cells (smooth-celled form),
x640 ; 8. upper laminal cells (papillose-celled form),
<640; 9. perichaetial leaf, x50; 10. capsule showing
line of dehiscence and spores, X20; 11. exothecial
cells and stomata at capsule base with spores, x90.
(I 3 & 9-1 1, Schelpe 7728; 4 & 6-7, Schelpe 7716a;
5, Schelpe 7780; 8, Goldblatt 639).
POTTIACEAE
209
papillae over lumen; marginal cells narrowly
rectangular or quadrate; basal cells larger,
rectangular, yellowish.
Autoicous. Antheridia in lower leaf axils;
perichaetia terminal, leaves undifferentiated.
Seta erect, 2-3 mm long, yellowish; capsule
cleistocarpic, exserted, reddish yellow, ovate
to cylindrical, 1,0-1, 8 mm high, long-
rostrate, 0,3 mm long, weakly oblique;
exothecial cells narrowly rectangular or some
cells quadrate or triangular, urn dividing at
middle when mature along weakly defined
dehiscence fine; stomata at base of urn,
phaneropore; calyptra large, cucullate, long-
rostrate, to 1 ,3 mm long, covering upper + of
capsule, becoming oblique with age; spores
subround to angular, 32-37 //m, verruculate.
Fig. 59: 1-11.
Endemic to Southern Africa, this species is
restricted to the shrublands of the western Cape
Province. Map 78.
Vouchers: Goldblatt 639; Schelpe 7716a, 7728,
7780; Tolken 1586a.
Pottia splachnoides is regarded as a highly variable
species encompassing a variety of leaf shapes and a
large variation in the degree of leaf cell papillosity,
and the size and shape of capsules. In view of similar
variability shown by the Australian species P. drum-
mondii (Wils.) Willis (Scott & Stone, 1976) it is prob-
able that the cleistocarpic species of Pottia present in
Australia [ P. drummondii], New Zealand [P. maritima
(R. Br. ter.) Broth.] and Southern Africa [P. splach-
noides] represent a single, widespread, Southern
Hemisphere taxon. The specimens recorded from
South Africa by Schelpe (1969) as P. maritima are
included here.
7. DESMATODON
Desmatodon Brid., Mant. Muse. 86 (1819); Grout, Moss FI. N. Amer. 1: 222 (1939); Saito in
Bull. nat. Sci. Mus., Tokyo 16: 66 (1973). Lectotype species: D. latifolius (Hedw.) Brid., vide
Venturi, Comment. Fauna FI. Veneto Trentino 1: 123 (1868).
Plants small, gregarious or loosely caespitose; terricolous or saxicolous. Stems 2-5 mm
tall, sparingly branched; with central strand. Leaves oval to lingulate; apex acute to obtuse;
margins plane to revolute, sometimes bordered, entire. Costa mucronate or percurrent,
strongly convex ventrally; in section guide cells 2-6, ventral cells large, surface cells slightly
larger than laminal cells, dorsal stereid band small. Laminal cells quadrate to hexagonal, with
numerous C-shaped papillae over lumen; marginal cells generally less papillose, sometimes
differentiated.
Autoicous or dioicous. Perichaetia terminal, leaves somewhat larger or undifferentiated.
Seta long, erect; capsule cylindrical, infrequently ovoid; peristome teeth divided into 2 long,
papillose filaments, fragile, frequently broken; operculum conical; calyptra cucullate; spores
round to reniform, smooth to punctate.
Desmatodon , a genus comprising 33 species, is best represented in temperate regions of the Northern
Hemisphere. Desmatodon convolutus is, however, found in semi-arid to temperate regions throughout the world.
In Southern Africa the genus is common in semi-arid shrublands of the Cape and South West Africa/Namibia,
but is also occasionally collected in grasslands of Natal and the Orange Free State.
1 Leaf margins revolute, marginal cells frequently smooth or almost so, not differentiated. ... 1. D. convolutus
1 Leaf margins plane, most leaves bordered by 3-5 rows of smooth, incrassate cells. ... 2 . D. longipedunculatus
1. Desmatodon convolutus {Brid.) Grout,
Moss FI. N. Amer. 1: 224 (1939); Flowers,
Moss. Utah 194 (1973). Type: Switzerland.
Trichostomum convolutum Brid., Muscol. Recent.
Suppl. 1: 232(1806).
Grimmia atrovirens J.E. Sm., Engl. Bot. 28: 2015
(1809). Tortula atrovirens (J.E. Sm.) Lindb. in Ofvers.
K. VetenskAkad. Forh. 21 : 236 (1864); Sim, Bryo. S.
Afr. 233 (1926).
Tortula recurvata Hook., Musci Exot. 2: 130
(1819); Dix. in Trans. R. Soc. S. Afr. 8: 194 (1920).
Barbula recurvata (Hook.) Schultz in Nova Acta
Acad. Leop. Carol. 11: 216 (1823). Desmatodon
recurvatus (Hook.) Mitt, in F. Mull., Fragm. Phyt.
Aust. Suppl. 11: 114 (1881); Catcheside, Moss.
South Austr. 148 (1980). Type: Cape, near Rogge-
veld, Burchell s.n. (BM, holo.l).
Tortula parvula Hook. & Grev. in Edinb. J. Sci.
1: 302 (1824). Barbula parvula (Hook. & Grev.)
210
POTTIACEAE
Spreng., Syst. Veg. 4: 179 (1827). Type: Cape of Good
Hope, Menzies s.n., 1791.
Didymodon capensis Spreng., Syst. Veg. 4: 179
(1827); C. Mull, in Linnaea 17: 598 (1843). Type:
Cape, Van Kamps Bay, Ecklon s.n., 1825.
Barbula circinalis C. Mull, in Linnaea 18: 703
(1845). Type: Cape, Swellendam, Ecklon s.n.
Tortula subtorquatifolia Dix. in K. norske Vidensk.
Selsk. Skr. 1932, 4: 8 (1932). Type: Cape, Vryburg,
H<f>eg 1 (BM, holo.!).
Plants small, gregarious or in loose
patches, dark green to blackish dry, green to
light green wet; terricolous or saxicolous.
Stems short, 2-5 mm high, sparingly
branched ; in section with central strand, inner
cortical cells lax, outer cortical cells in 1-3
rows, smaller, reddish. Leaves incurled,
spirally twisted or contorted dry, wide-
spreading wet; shape extremely variable, oval
to oblong on short-leaved plants or oblong to
lingulate on long-leaved plants; apex acute to
obtuse; margins narrowly revolute, entire.
Costa percurrent, strongly convex ventrally in
upper leaf; ventral superficial cells quadrate,
papillose, dorsal superficial cells elongate,
strongly incrassate, smooth; in section oval,
guide cells 4-6, ventral cells in 1-2 layers,
surface cells enlarged, frequently vertically
elongated, papillose, dorsal stereid band 4-6
cells thick, dorsal surface cells substereids.
Upper laminal cells quadrate to hexagonal,
with 4-6 C-shaped papillae over lumen; cells
of revolute margins smooth or nearly so,
rarely all laminal cells only weakly papillose;
basal cells rectangular, lax, smooth.
Autoicous. Perichaetia terminal, leaves
undifferentiated. Seta 6-15 mm long, yellow
to reddish; capsule ovoid to cylindrical,
1,0-2, 5 mm long, reddish brown; peristome
teeth divided into 2 papillose filaments above
short basal membrane, 0,3-0, 5 mm high,
yellowish, frequently irregular, broken; oper-
culum conical to obliquely rostrate, 1 mm
long; spores roundish, 17-25 pm, finely
punctate. Fig. 60: 1-10.
Desmatodon convolutus is found in Europe, the
Middle East, Africa, Micronesia, Australia, New
Zealand, North America and southern South America.
In Southern Africa, the species is common in arid and
semi-arid shrublands throughout the Cape Province.
It is also frequently collected in the Orange Free State,
Lesotho and southern South West Africa/Namibia,
and occasionally in the Transvaal and Natal. Map 79.
Vouchers: Emdon 203a; Lavranos 15470; Mac-
Donald 11 Mag ill 4177, 5846; Schelpe 7783, 7821 ;
Smook & Harding 799; Volk 6642.
Map 79. — • Desmatodon convolutus
x Desmatodon longipedunculatus
Sometimes troublesome in its many forms, D.
convolutus can generally be recognized by the enlarged,
upper ventral costal cells. Some of the smaller plants
with rounded leaves may be confused with Pottia.
These smaller forms may also occasionally have
rudimentary peristomes that make identification even
more difficult. The presence of a central strand in the
stem and the more highly developed costal anatomy,
will separate Desmatodon from Pottia.
2. Desmatodon longipedunculatus (C.
Mull.) Magill, comb. nov. Type: Cape,
Swellendam, Pappe s.n., 1838 (H, iso.!).
Barbula longipedunculata C. Mull.,Syn. Muse. 1 : 630
(1849). Tortula longipedunculata (C. Mull.) Broth, in
Naturl. PflFam. 1 : 432 (1902); Sim, Bryo. S. Afr. 230
(1926).
Barbula desertaC. Miill. in Hedwigia 38: 108 (1899).
Tortula deserta{C. Miill.) Broth, in Naturl. PflFam. 1:
430 (1902). Type: Cape, Cape Town, Rehmann ?96,
1875.
Plants small, caespitose, greyish green to
blackish green dry, yellowish green wet,
reddish brown below; terricolous. Stems 2-4
mm tall, unbranched; in section with large
central strand, cortical cells lax, single outer
row of cells smaller, incrassate, red-brown.
Leaves appressed, spirally twisted dry, patent
wet; oval to lingulate, 1,0-2, 6 mm long;
apex obtuse; margins plane, entire, marginal
cells differentiated, forming unistratose bor-
der 3-5 cells wide. Costa mucronate; ven-
tral superficial cells quadrate, papillose,
dorsal superficial cells rectangular, thickened,
with scattered, low, blunt papillae; in section
with 2-4 guide cells, ventral cells in 1-2 rows,
POTTIACEAE
211
Fig. 60. — Desmatodon convolutus (1-10): 1.
habit, x 1 ; 2. habit, wet and dry, x 10; 3. stem in cross
section, x90; 4. leaves, x25; 5. leaf in distal cross
section, x220; 6. lower leaf cells, xl70; 7. cells of
lamina (dorsal surface), xl70; 8. leaf apex (papillae
partly shown), xl70; 9. perichaetial leaves, x25;
10. part of capsule mouth with peristome teeth and
spores, x 100. D. longipedunculatus (11-20): 11.
habit, x 1; 12. habit, x 10; 13. stem in cross section,
x90; 14. lower leaf, x25; 15. upper leaf, x25; 16.
leaf in distal cross section, x220; 17. basal leaf cells
at margin, x 170; 18. upper laminal cells at right
margin, x375; 19. leaf apex, xl70; 20. part of
capsule mouth with peristome teeth, x 100. (1-10,
Schelpe 7823; 11-20, Magill & Schelpe 3919).
212
POTTIACEAE
large, surface cells papillose, dorsal stereid
band 4-5 cells thick, dorsal surface cells
substereids, 1-2 lateral surface cells similar to
ventral surface cells. Upper laminal cells
quadrate to hexagonal, some transversely
rectangular, with 4-5 low, C-shaped papillae
over lumen; marginal cells quadrate to
rectangular, some transversely rectangular,
incrassate, smooth to weakly papillose,
generally forming distinct border above base;
basal cells quadrate to rectangular, smooth,
smaller toward margin.
Dioicous. Perichaetia terminal, leaves
undifferentiated. Seta 10-12 mm long, yellow
to red; capsule erect, cylindrical, 2, 0-2, 5 mm
long, red-brown; peristome teeth divided to
base into 2 long papillose filaments, 0,8 mm
long, spirally twisted one turn; operculum
conical, 1 ,0 mm long; spores round, to
reniform, 10-12 pm, smooth. Fig. 60: 11-20.
Originally described from the fynbos biome of the
southwestern Cape, D. longipedunculatus is also found
in the dwarf succulent shrublands of the northwestern
Cape and the Khomas Highland Savanna of South
West Africa/Namibia. The species was recently
reported from Kenya by Bizot, Poes & Sharp (1979).
Map 79.
Vouchers: Magill & Schelpe 3884a; Tolken 5684;
Volk 00950.
Although not always pronounced, the plane,
bordered leaf margins will separate this species from
D. convolutus. In section the cells along the le;.f
margins are slightly larger, incrassate, and nearly
smooth. This weak border is generally obvious in
surface view, but occasionally the cells are papillose
and the border is only detectable in cross section.
8. TORTULA
Tortula Hedw., Sp. Muse. 122 (1801), nom. cons.; Broth, in Naturl. PflFam. 10: 295 (1924);
Steere in Grout, Moss FI. N. Amer. 1 : 228 (1939); Sim, p.p., Bryo. S. Afr. 224 (1926). Type
species: T. subulata Hedw.
Plants small to large, forming large tufts, dark green to yellowish or brownish green:
corticolous, saxicolous or terricolous. Stems frequently branched; generally with central
strand. Leaves appressed, twisted or contorted dry, widespreading wet; piliferous, triangular
to spathulate; apex acute to emarginate; margins plane to revolute. Costa mucronate to long-
excurrent, smooth to denticulate, reddish yellow to hyaline; in section without ventral stereid
band, dorsal stereid band large, prominent. Laminal cells quadrate to polygonal, with nume-
rous C- or O-shaped papillae, rarely with single spinose papilla per cell; basal cells generally
larger, quadrate to rectangular, smooth, hyaline. Propagulae leaf-like, produced in stem apex;
gemmae round, multicellular, produced on upper ventral costal surface or on stem tomentum.
Autoicous, synoicous or dioicous. Perichaetial leaves undifferentiated. Capsule long-
exserted, cylindrical; peristome teeth divided into 32 filaments above basal membrane, spirally
twisted, papillose; operculum conical; calyptra cucullate; spores round, finely granular,
punctate or smooth.
Tortula, with 250 species, is found throughout the world in a great variety of habitats. In Southern Africa
the genus is frequently collected in the Cape and Natal; however, specimens are found throughout the
Flora area.
1 Costa mucronate:
2 Leaves fragile, especially below; sterile plants without propagulae; capsule mouth with several rows of
smaller cells 3. T. schmidii
2 Leaves not fragile; leaf-like propagulae produced on short stalks at apex or in leaf axils; capsule mouth
with 2-3 rows of smaller cells 4. T. ammonsiana
1 Costa apiculate to long-excurrent:
3 Costa apiculate to short-excurrent:
4 Leaves ovate-acuminate to triangular; margins spirally revolute; marginal cells enlarged; gemmae
not known 1 . T. porphyreoneura
4 Leaves broadly spathulate; margins plane; marginal cells undifferentiated; gemmae produced on
ventral costal surface 6. T. papillosa
POTTIACEAE
213
3 Costa long-excurrent:
5 Awn smooth or almost so:
6 Plants corticolous; leaf margins plane; propagulae produced at stem apex 5. T. pagorum
6 Plants terricolous; leaf margins revolute; without propagulae 2.T. muralis
5 Awn denticulate:
7 Plants synoicous; leaves widespreading when wet; awn weakly denticulate; apex obtusely acute
to retuse; stem with small central strand 7. T. princeps
7 Plants dioicous; leaves recurved to squarrose; awn roughly denticulate; apex emarginate; stem
without central strand 8. T. ruralis
1 . Tortula porphyreoneura (C. Miill.)
Townsend in J. Bryol. 10: 576 (1979). Type:
Ethiopia, Abita Keren, Beccari s.n., 1870 (FI).
Barbula porphyreoneura C. Mull, ex Vent, in Nuovo
G. bot. ital. 4: 13 (1872).
Tortula torquatifolia (Geh.) Dix. in K. norske
Vidensk. Selsk. Skr. 1932 (4): 8 (1932); Hilpert in
Beih. bot. Zbl. 50: 626 (1933); fide Townsend in J.
Bryol. 10: 576 (1979). Barbula torquatifolia Geh. in
Bull. Herb. Boissier 4: 410 (1896); Broth, in Natiirl.
PflFam. 1: 411 (1902); 10: 280 (1924); Sim, Bryo. S.
Afr. 235 (1926). Type: South West Africa/Namibia,
Oshando, H. Schinz s.n., 1886 (G, holo.!; Z).
Plants small, in loose tufts, green; terri-
colous. Stems 5-10 mm tall, lower parts
frequently buried, unbranched; in section
with central strand, inner cortical cells large,
lax, outer cortical cells in 1-3 rows, smaller,
incrassate, red-brown. Leaves imbricate, con-
torted or spirally twisted dry, spreading wet;
ovate-acuminate to triangular, 1,5-2, 5 mm
long; apex acute to obtuse; margins spirally
revolute, entire; marginal cells differentiated
into specialized photosynthetic region; in
section 4-5 cells enclosed by spiralling leaf,
enlarged, thin-walled. Costa apiculate to
Map 80. — • Tortula porphyreoneura
X Tortula princeps
short-excurrent as smooth, yellow awn, 35-40
p m wide at apex; ventral superficial cells
quadrate, papillose, dorsal superficial cells
rectangular, incrassate, smooth; in section
guide cells 2, ventral cells 2, papillose, dorsal
stereid band 5-8 cells thick, dorsal surface
cells frequently substereids. Upper laminal
cells quadrate to hexagonal, juxtacostally
with 4-6 C- or O-shaped papillae over lumen,
cells of revolute portion smooth or ventrally
papillose; marginal cells enlarged, lax; basal
cells quadrate, smooth.
Dioicous. Perichaetia terminal, leaves
undifferentiated. Seta 5-7 mm long, yellowish
red, becoming red with age; capsule cylindri-
cal, 1,5-1, 8 mm long, red-brown; peristome
short, 0,6 mm high, teeth linear above short
basal membrane, spirally twisted 1-2 times,
papillose; operculum rostrate, 0,8 mm long;
spores round, 10-11 pm, smooth or weakly
punctate. Fig. 61: 1-9.
Tortula porphyreoneura is known from Arabia
and eastern and Southern Africa. In the Flora area the
species is infrequently collected around rock outcrops
in grasslands in northern South West Africa/Namibia,
Botswana, and the northern and eastern Transvaal.
Map 80.
Vouchers: Hardy 5173; Leistner et al. 72; Magill
4761, 4954; Tolken 5559.
The macroscopic resemblance of this species to
Barbula is superseded by the Tortula-Uke peristome
and costal anatomy. The differentiated marginal leaf
cells would place this species in Pseudocrossidium sensu
Zander (1979); see note under Barbula acutata.
2. Tortula muralis Hedw., Sp. Muse. 123
(1801); Broth, in Natiirl. PflFam. 10: 297
(1924); Sim, Bryo. S. Afr. 230 (1926). Type:
Europe.
Barbula chrysoblasta C. Miill. in Hedwigia 38: 104
(1899). Type: Cape, Uitenhage, Breutel s.n.
Barbula afroinermis C. Miill. in Hedwigia 38: 104
(1899). Type: Cape, Cape Town, Naumann s.n., 1874
(BM, iso.!).
Plants small, in dense tufts, yellow-green
to dark green; terricolous. Stems 4— 8(— 12)
214
POTTIACEAE
PoTTIACEAE
215
mm tall, branched below; in section with
central strand, cortical cells larger toward
margin, reddish. Leaves erect, contorted or
spirally twisted dry, spreading wet; oblong-
spathulate to Ungulate, 2-3 mm long; apex
obtuse, retuse or emarginate, often asym-
metrical; margins revolute to apex, entire.
Costa excurrent, 1-2 mm long, smooth,
hyaline; ventral superficial cells quadrate to
rectangular, thin-walled, papillose, dorsal
superficial cells elongate-rectangular, incras-
sate, smooth ; in section guide cells 2-4, ventral
cells 4-8, in single or double row, surface cells
papillose, dorsal stereid band 6-8 cells thick,
dorsal surface cells undifferentiated, smooth.
Upper laminal cells quadrate to hexagonal,
densely papillose with 6-8 C-shaped papillae
over lumen; basal cells rectangular, hyaline,
smooth.
Autoicous. Perichaetial leaves undifferen-
tiated. Seta 10-20 mm long, yellowish,
becoming reddish to red-brown with age;
capsule cylindrical, 2, 5-3,0 mm long, red-
brown, peristome 0,6 mm high, teeth linear,
above short basal membrane, papillose,
spirally twisted 1-2 times; operculum
conical, 1 mm long; spores round, 9—12 pm,
weakly papillose. Fig. 61: 17-23.
Tortula muralis is almost cosmopilitan in its
distribution. In Southern Africa the species is fre-
quently collected in the southwestern Cape, especially
in and around Cape Town. A few specimens are also
known from southern South West Africa/Namibia,
the western, southern and eastern Cape and Natal.
The species is commonly collected on old rock walls
or other cement structures. Map 81.
Vouchers: Cholnoky 860a; Ellis 3102; Esterhuysen
18880; Garside 6569; Magill 6246, 6330; Sim 9312.
The plants are considerably smaller than the other
species of Tortula in Southern Africa. The smooth,
incrassate, ± rectangular cells at the leaf apex, useful
for identification in other regions, are not always
present in the Southern African plants. This appears
however, to be the only difference between Southern
and Northern Hemisphere plants. The presence of
T. muralis , primarily on man-made structures,
indicates its introduction by man.
Map 81. — • Tortula ammonsiana
x Tortula muralis
3. Tortula schmidii (C. Mull.) Broth, in
Natiirl. PflFam. 1: 434 (1902); Townsend in
J. Bryol. 10: 129 (1978). Type: India.
Barbula schmidii C. Mull, in Bot. Ztg 11: 58 (1853).
Barbula erubescens C. Miill. in Nuovo G. bot. ital.
4: 14 (1872). Tortula erubescens (C. Miill.) Broth, in
Natiirl. PflFam. 1: 434 (1902), horn, illeg.; Sim, Bryo.
S. Afr. 227 (1926). Type: Ethiopia, Bogos, Keren,
Beccari s.n. (FI).
Barbula hildebrandtii C. Miill. in Linnaea 40: 294
(1876). Tortula hildebrandtii (C. Miill.) Broth, in
Natiirl. PflFam. 1: 434 (1902). Type: Somalia,
Hildebrandt sub 1 502.
Barbula br achy aichmeC. Miill. in Hedwigia 38: 102
(1899). Tortula brachyaichme (C. Mull.) Broth, in
Natiirl. PflFam. 1: 434 (1902). Type: Cape, Cape
Town, Rehmann s.n. (BM!).
Barbula exesaC. Miill. in Hedwigia 38: 103 (1899).
Tortula exesa (C. Miill.) Broth, in Natiirl. PflFam. 1 :
434 (1902). Type: Cape, Somerset East, Boschberg,
MacOwan s.n., 1883.
Barbula macowaniana C. Miill. in Hedwigia 38: 103
(1899). Tortula macowaniana (C. Miill.) Broth, in
Natiirl. PflFam. 1: 434 (1902). Type: Cape, Somerset
East, MacOwan s.n., 1878 (GRA!).
Barbula oranica C. Miill. in Hedwigia 38: 103
(1899). Tortula oranica (C. Miill.) Broth, in Natiirl.
PflFam. 1: 434 (1902). Type: Orange Free State,
Bethlehem, Rehmann 126 (PRE!).
Fig. 61. — Tortula porphyreoneura (1-9): 1. habit, xl; 2. habit, xlO; 3. stem in cross section, xlOO; 4.
leaves, x 35 ; 5. leaf in cross section, x 280; 6. basal leaf cells, x 170; 7. upper laminal cells, x 640; 8. leaf apex,
x 170; 9. part of capsule mouth with peristome teeth and spores, x 75. T. schmidii (10—16) : 10. habit, x 1 ; 11.
habit, x 10; 12. stem in cross section, x 100; 13. leaves, x 20; 14. leaf in cross section, x 175; 15. upper laminal
cells, x425; 16. parts of capsule mouth and peristome, x75. T. muralis (17-23): 17. habit, x 1 ; 18. habit, x 10;
19. leaves, x 20; 20. leaf in cross section, x 175: 21. upper laminal cells, x480; 22. cells at leaf apex (left side),
x 170; 23. deoperculate capsule, x 10; 24. capsule mouth and peristome, x 100. (1-9, Leistner et al. 72; 10-15,
Sim 8669; 16-18, Magill 6246; 19-24, Sim 9202).
216
POTTIACEAE
Barbula brevimucronata C. Miill. in Hedwigia 38 :
104(1899). Tortula brevimucronata (C. Miill.) Broth, in
Natiirl. PflFam. 1: 434 (1902). Type: Transvaal,
Spitzkop, near Lydenburg, Wilms s.n., 1888 (G,
holo.!).
Tortula brevitubulosa Broth, in Denkschr. Akad.
Wiss., Wien. Math.-nat. Kl. 88: 736 (1913). Type:
Natal, Van Reenens Pass, Schwarzerberg, Brunthaler
s.n., 4/12/1909 (H-BR, holo.!).
Tortula irregularis Sim, Bryo. S. Afr. 231 (1926).
Type: Natal, Edendale Falls, Sim 10064 (PRE, holo.!).
Plants medium-sized, forming dense
tufts, dark green to olive-green; corticolous,
less frequently saxicolous or terricolous.
Stems 10-15 mm high, occasionally branched,
tomentose below; in section with small central
strand, inner cortical cells large, thin-walled,
outer cortical cells in 3-4 rows, smaller,
incrassate, yellow-red. Leaves appressed,
contorted dry, widespreading to recurved wet,
fragile, lamina frequently broken or absent in
lower stem; lingulate to spathulate, 2, 5-3,0
mm long; apex obtuse; margins recurved
below, plane above, irregularly and weakly
lobed in upper leaf. Costa mucronate to
apiculate; ventral superficial cells quadrate,
thin-walled, papillose, dorsal superficial cells
elongate, strongly incrassate, denticulate
above ; in section with 2-4 large guide cells,
ventral cells incrassate, in 2 layers, ventral
surface cells papillose, dorsal stereid band
large, 6-8 cells thick, dorsal surface cells not
differentiated. Upper laminal cells quadrate
to hexagonal, with 4-6 C-shaped papillae over
lumen; basal cells rectangular to quadrate,
smooth; upper basal marginal cells quadrate.
Sporophytes not known in Southern
Africa. Fig. 61 : 10-16.
Known from India and eastern Africa south to
the Cape, the species is common in forest and wood-
land. Within the Flora area, T. schmidii is most fre-
quently collected on trees in the forests of the eastern
Transvaal, Swaziland, Natal, Transkei and the eastern
and southwestern Cape, or on soil in the western Cape
and rocks in the high grasslands of Lesotho and at
Mont-aux-Sources in the Orange Free State. Map 82.
Vouchers: Crosby & Crosby 8160; Ellis 3095;
Magi 1 1 4613a, 4854, 5670; Schelpe 7549; Smook 1104.
The plants are identified by the fragile leaves with
lamina frequently completely broken away on the
lower stem leaves. The species is similar to T. ammon-
siana, but that species produces abundant stalked
propagulae, has non-fragile leaves and only 2-3 rows
of smaller cells at the capsule mouth. The eastern
African capsules of T. schmidii have 5-6 rows of
smaller cells at capsule mouth (cf. Townsend, 1978).
4. Tortula ammonsiana Crum & Anderson
in Bryologist 82: 469 (1979). Type: North
America, West Virginia, Anderson 21897
(DUKE, holo.; MICH; TENN; WVA).
Plants medium-sized, forming loose tufts,
green to dark green above, light brown below;
saxicolous or corticolous. Stems 10-15 mm
high, branching; in section with large central
strand, inner cortical cells large, thin-walled,
outer cortical cells in 1-2 rows, small, incras-
sate, reddish. Leaves imbricate, contorted dry,
widespreading wet; lingulate to subspathu-
late, 2, 5-4, 5 mm long; apex rounded,
margins entire, plane or reflexed at mid-leaf.
Costa mucronate; ventral superficial cells
quadrate or infrequently rectangular, thin-
walled, papillose, dorsal superficial cells
elongate, strongly incrassate, smooth; in
section guide cells 2, ventral cells in two
layers, surface cells papillose, dorsal stereid
band 6-8 cells thick, dorsal surface cells not
differentiated. Upper laminal cells quadrate
to hexagonal, with 4-6 C-shaped papillae over
lumen; basal cells rectangular, hyaline,
smooth ; upper basal marginal cells quadrate.
Propagulae leaf-like, in clusters on short stalks
at apex; elliptical; costa present, ending as
smooth, yellow apiculate cell; body cells
hexagonal, papillose.
Perichaetia terminal, leaves slightly smal-
ler. Seta erect, 7-9 mm long, reddish yellow;
capsule cylindrical, 2,0-2, 5 mm long,
reddish yellow, exothecial cells rectangular,
1-3 rows at mouth smaller, quadrate; peri-
stome fragile, yellowish to reddish yellow,
teeth weakly twisted, filamentous, ornately
papillose above short, erectly tasselated basal
membrane; mature operculum not seen,
calyptra cucullate, 4 mm long; spores round,
15 pm, papillose. Fig. 62: 1-11.
This species was very recently described from the
eastern United States. In Southern Africa the species
is found in forests or grasslands of the eastern Trans-
vaal, Orange Free State, Lesotho, Natal and southern
and southwestern Cape. Map 81.
Vouchers: Magill 4595, 4860, 6268; Van Rooy
404, 572.
This species is recognized by its mucronate
leaves and the production of numerous, stalked,
leaf-like propagulae. The Southern African specimens
differ from the North American plants in the absence
of the minute apical teeth only. The species is related
to T. schmidii and T. pagorum ; differences are discussed
under those species.
POTTIACEAE
217
5. Tortula pagorum ( Milde ) De Not. in
Atti Univ. Genova 1: 542 (1869); Broth, in
Natiirl. PflFam. 10: 300 (1924); Steere in
Grout, Moss FI. N. Amer. 1: 239 (1939).
Type : Switzerland, Milde s.n.
Barbula pagorum Mild, in Bot. Ztg 20 : 469 (1862).
Plants small, in loose tufts, dark to olive-
green; corticolous. Stems 2-4 mm high,
unbranched ; in section central strand present,
inner cortical cells large, thin-walled, outer
cortical cells in 1-2 rows, smaller, red-brown.
Leaves appressed, incurved dry, widely
spreading wet, comose ; spathulate to broadly
lingulate, 2 mm long; apex emarginate to
retuse; margins plane, entire. Costa excurrent
as long, smooth, hyaline awn; ventral super-
ficial cells quadrate, thin-walled, papillose,
dorsal superficial cells linear, strongly incras-
sate, smooth; in section with 2 guide cells,
ventral cells in one layer, thin-walled, papil-
lose, dorsal stereid band large, 7-8 cells thick.
Laminal cells quadrate to hexagonal, with 4-8
C-shaped papillae over lumen; basal cells
larger, quadrate to shortly rectangular,
Fig. 62. — Tortula ammonsiana: 1. habit, plant
with apical propagulae, xl; 2. habit, plant with
sporophyte, x 1 ; 3. habit, plant with apical pro-
pagulae, xlO; 4. stem in cross section, x80; 5.
leaves, x25; 6. leaf in cross section, xl70; 7. basal
leaf cells at left margin, x 170; 8. upper laminal cells
(papillae partly shown), x435; 9. leaf apex (papillae
partly shown), x 170; 10. propagulum, x 140; 11.
capsule mouth with peristome, x80. (1-11, Crosby &
Crosby 8160).
218
POTTIACEAE
smooth. Propagulae leaf-like, in large numbers
at stem apex; elliptical; ecostate; apical cell
elongate, pellucid, body cells hexagonal to
subhexagonal, papillose, chlorophyllous.
Sporophyte not known in Africa, but
described from Australia (Stone, 1971) as:
Perichaetial leaves little differentiated. Seta
5-9 mm long, yellow above, reddish below;
capsule short-cylindrical 2, 0-2, 4 mm long,
slightly curved, brownish; peristome to 1,0
mm high, teeth linear, papillose, twisted once
counter-clockwise, above a short basal mem-
brane, 0,3 mm high; operculum narrowly
conical, 1 ,5-1,7 mm long; calyptra cucullate,
2, 5-3,0 mm long; spores 8—10 pm, slightly
papillose, greenish brown. Fig. 63: 1-8.
Tortula pagorum is known from the Americas,
Europe, Asia, Africa and Australia, generally in
association with human habitation. In Southern
Africa the species has been infrequently collected in
the northern, central and eastern Transvaal, Swazi-
land, southern Natal and the eastern, central, southern
and southwestern Cape. The species is most commonly
found on bark of small trees or shrubs, in dry wood-
land communities. Map 83.
Map 83. — • Tortula pagorum
X Tortula papillosa
Vouchers: Magill 3198, 3514, 4921a, 6142; Sim
10083; Smook 990.
Tortula pagorum is recognized by its long-
excurrent costa and the production of numerous
propagulae at the stem apex. The species is related to
T. ammonsiana, the only other Southern African
species of Tortula that produces propagulae, but
differs in the excurrent leaf costa and absence of
costae in the propagulae that are sessile at the apex.
6. Tortula papillosa Wils. ex Spruce in J
Bot., Lond. 4: 193 (1845); Broth, in Natiirl.
PflFam. 10: 300 (1924); Sim, Bryo. S. Afr. 226
(1926). Type: England.
Barbula reticularia C. Mull, in Hedwigia 38: 101
(1899). Tortula reticularia (C. Mull.) Broth, in Natiirl.
PflFam. 1: 434 (1902). Type: Cape, Cape Town,
Rehmann 106 (PRE!).
Plants small, gregarious or in small tufts,
green to brown-green; corticolous. Stems 2-8
mm high, infrequently branched; in section
with central strand, cortical cells large,
becoming smaller toward margin, yellow-
brown. Leaves imbricate with incurved apices
dry, widespreading wet; broadly spathulate,
2,0-2, 5 mm long; apex obtusely acute to
rounded ; margins plane, entire. Costa spinose
dorsally, gemmiferous ventrally, excurrent as
short, yellowish, weakly toothed awn; ventral
superficial cells producing numerous gemmae,
dorsal superficial cells elongate, incrassate,
sparsely papillose; in section with 2 guide
cells, ventral cells 4, in single row, thin-walled,
producing gemmae, dorsal stereid band 3-6
cells thick. Laminal cells hexagonal to poly-
gonal, 20-25 pm, smooth ventrally, dorsally
with l(-2) C- or O-shaped, spinose papillae
over lumen; basal cells rectangular, smooth.
Gemmae round, multicellular, 50-80 pm,
green.
Sporophyte not known in Southern
Africa. Sporophytes from Australia, illus-
trated by Scott & Stone (1976), show differen-
tiated perichaetial leaves that sheath the seta.
The capsule is similar to that of T. pagorum,
but the seta is shorter and twisted to the right.
Fig. 63:9-16.
Tortula papillosa is known from Europe and
North America, southern South America, South
Africa, Australia and New Zealand. The species is
rare in Southern Africa. It is collected on bark of trees
in dry woodlands and forests of Natal, and the eastern
and southwestern Cape. Map 83.
Vouchers: Ellis 3092; Sim 9570, 10082.
This species is separated from the other Southern
African species of Tortula by the single spinose papilla
on the dorsal surface of leaf cells and in the production
of multicellular gemmae on the upper ventral costal
surface.
7. Tortula princeps De Not. in Memorie
Accad. Sci. Torino 40: 288 (1838); Broth, in
Natiirl. PflFam. 1:435(1902); 10:302(1924);
Grout, Moss FI. N. Amer. 1: 244 (1939).
Type: Sardinia, Rabenhorst s.n., 1847.
POTTIACEAE
219
Barbula muelleri B.S.G., Bryol. Eur. 2: 106 (1842),
nom. illeg. Tortula muelleri Hook. f. & Wils., FI.
Antarct. 1 : 103 (1847), nom. illeg.; Sim, Bryo. S. Afr.
225 (1926).
Barbula leucostega C. Mull., Syn. Muse. 1 : 641
(1847). Tortula leucostega (C. Miill.) Broth, in Natiirl.
PflFam. 1: 435 (1902). Type: Cape, Swellendam,
Ecklon s.n., 1828.
Plants large, in dense tufts, green to
yellow-brown; terricolous. Stems 10-30 mm
high, branching, tomentose below; in section
with central strand, cortical cells lax,
becoming smaller toward margin, yellowish
red. Leaves appressed, keeled, little contorted
dry, erect to widespreading wet; broadly
lingulate to spathulate, 3-4 mm long; apex
obtusely acute to retuse; margins narrowly
revolute below, plane above. Costa long-
excurrent as hyaline awn, 1-2 mm long,
weakly denticulate; ventral superficial cells
quadrate, thin-walled, papillose, dorsal super-
ficial cells elongate, strongly incrassate,
smooth; in section guide cells 2, ventral cells
in 1-2 layers, surface cells papillose, dorsal
stereid band large, 4-5 cells thick, dorsal
surface cells undifferentiated, smooth. Lami-
nal cells quadrate to hexagonal, densely
papillose, with 4-8 C- or O-shaped papillae
over both surfaces; basal cells large, in well
defined group, rectangular, hyaline, smooth.
Synoicous. Perichaetia terminal, leaves
undifferentiated. Seta 15-20 mm long, red-
brown; capsule cylindrical, 3-4 mm long,
red-brown ; peristome reddish yellow to white,
0,8-1 ,0 mm high, teeth papillose, filamentous
above a high, obliquely tessellated basal
membrane; operculum conical, 1,0-1, 5 mm
long; spores round, 10-15 //m, almost smooth.
Fig. 63: 17-23.
Tortula princeps is a very widespread species in
temperate regions of both hemispheres. In Southern
Africa T. princeps has been occasionally collected in
shrublands of the western Cape Province. Map 80.
Vouchers: Gar side 6708 ;Lavranos 15202a; Wager
632.
The Southern African specimens are not always
clearly separated from T. ruralis. In the absence of the
synoicous inflorescence, a large number of characters
must be assessed to separate the species. Many of the
recent collections are satisfactorily placed, but a few
sterile specimens have been collected that appear to
be intermediate between the two species.
8. Tortula ruralis ( Hedw .) Gaertn., Meyer
& Scherb. in Oek, Techn. FI. Wetterau 3: 91
(1802); Broth, in Natiirl. PflFam. 1: 434
(1902); 10: 301 (1924); Sim, Bryo. S. Afr. 226
(1926). Type: Germany.
Barbula ruralis Hedw., Sp. Muse. 121 (1801).
Barbula afroruralis C. Miill. in Hedwigia 38: 101
(1899). Tortula afroruralis (C. Miill.) Broth, in Natiirl.
PflFam. 1: 435 (1902). Type: Cape, Stinkwater,
Rehmann 114 (PRE!; NH!).
Barbula erythroneura C. Miill. in Hedwigia 38: 102
(1899). Tortula erythroneura (C. Miill.) Broth, in
Natiirl. PflFam. 1: 435 (1902). Type: Cape, Groene-
kloof, Breutel s.n., 1862.
Tortula trachyneura Dix. in Trans. R. Soc. S. Afr.
8: 195 (1920). Type: Cape, Tjumie, D. Henderson 330
(BM, holo.!).
Plants large, in loose tufts, green to
brownish green; terricolous or saxicolous.
Stems 5-15 mm tall, branching, tomentose
below, tomentum rarely gemmiferous; in
section without obvious central strand, inner
cortical cells large, thin-walled, outer cortical
cells in 1-3 rows, smaller, incrassate, reddish.
Leaves appressed, rarely bistratose, mostly
contorted dry, recurved to squarrose wet;
oblong-spathulate, 2-3 mm long; apex emar-
ginate or truncate, frequently asymmetrical;
margins revolute to upper leaf, frequently to
apex ; base appressed. Costa long-excurrent as
hyaline, dentate awn, 2-3 mm long; ventral
superficial cells quadrate to short-rectangular,
thin-walled, papillose, dorsal superficial cells
elongate, incrassate, smooth to rough or
spinose; in section guide cells 2, ventral cells
in 1-2 layers, large ventral surface cells
papillose, dorsal stereid band 4-5 cells thick,
reddish, dorsal surface cells generally hyaline,
smooth, papillose or spinose. Laminal cells
hexagonal, densely papillose, with 4-10
220
POTTIACEAE
POTTIACEAE
221
C-shaped papillae over lumen; basal cells
large, in well defined group, rectangular,
hyaline, smooth; basal marginal cells qua-
drate, chlorophyllous, papillose. Gemmae
round, multicellular, 50-60 /am, green, rarely
on tomentum of lower stem or ventral costal
surface.
Dioicous. Perichaetia terminal, leaves
undifferentiated. Seta 10 mm long, reddish
brown; capsule cylindrical, 2-3 mm long,
brownish; peristome reddish to white, 1,5
mm high, basal membrane long, obliquely
tessellated, teeth long-filamentous, papillose;
operculum conical, 1 mm long; spores 12-15
//m, finely granular. Fig. 63 : 24-32.
Widespread in Europe, Asia and North America,
T. ruralis is less common in Southern Africa and
Patagonia. The species, growing on soil and rock in
shrubland or grassland, has been collected in a few
locations in the Transvaal, Orange Free State,
northern Cape, Lesotho and Transkei. It is more
common in the southern and western Cape. Map
84.
Vouchers: Esterhuysen 20600; Hardy 4315; Lyle
7078; Magill 3293, 4525, 6120; Schelpe 7816.
The large, squarrose leaves with strongly spinose
awn and the dioicous condition characterize this
species. The strongly spinose, dorsal costa ( T . tra-
chyneura Dix.) is seen as only a modification, as are
the bistratose leaves found in several arid region
collections. Sporophyte production and occurrence of
rhizoid or costal gemmae are rare in this species in
Southern Africa.
9. HUSN OTIELL A
Husnotiella Card, in Rev. Bryol. 36: 71 (1909); Broth, in Natiirl. PflFam. 10: 256 (1924). Type
species : H. revoluta Card.
Plants small to very small, in cushions, green; on soil or soil over rock. Stems erect,
branching; with central strand. Leaves appressed dry, erect to recurved wet; lingulate to
oval-lanceolate ; apex broadly acute to rounded ; margins recurved, entire to crenulate. Costa
ending below apex; in section round, cells scarcely differentiated, incrassate. Laminal cells
rounded, quadrate to hexagonal, incrassate, smooth or papillose; basal cells rectangular,
thin-walled, smooth.
Apparently dioicous. Perichaetial leaves generally distinct. Seta erect, 5-9 mm long,
capsule cylindrical to ovate, smooth or weakly plicate, gymnostomous; operculum rostrate;
calyptra cucullate; spores round, smooth to granulate.
The genus Husnotiella contains approximately seven species. With the exception of the two Southern
African species, the genus is strictly American in its distribution.
1 Leaf cells smooth 1. H. latifolia
1 Leaf cells papillose 2. H. plicata
1. Husnotiella latifolia (Dix.) Zander &
Magill in Mem. bot. Surv. S. Afr. 43 : 7 (1979).
Type; Zimbabwe, Victoria Falls, Sim 8931
(BM, holo.l; PRE1).
Gyroweisia latifolia Dix. in S. Afr. J. Sci. 18: 309
(1922); Sim, Bryo. S. Afr. 251 (1926).
Gyroweisia amplexicaulis Sim, Bryo. S. Afr. 250
(1926). Type: Natal, Van Reenen, Wager 414 (PRE,
holo.l).
Fig. 63. — Tortula pagorum (1-8): 1. habit, x 1 ; 2. habit, plant with apical propagulae, x 10; 3. leaf, x 20;
4. leaf in cross section, x200 ; 5. basal leaf cells (upper right side), xl70; 6. upper laminal cells, x640; 7.
leaf apex (left side), x 170; 8. propagulum, x 140. T. papillosa (9—16) : 9. habit, x 1 ; 10. habit, x 10; 11. stem in
cross section, x85; 12. leaf, x20; 13. leaf in distal cross section, xl70; 14. upper laminal cells, x640; 15.
cells at leaf apex, x 170; 16. gemmae, x 200. T. princeps (17-23): 17. habit, x 1 ; 18. leaf, x 20; 19. leaf in cross
section, x200 ; 20. cells at basal leaf margin, x 170; 21. upper laminal cells, x640; 22. part of capsule mouth
with peristome teeth, x30; 23. young sporophyte at stem apex with surrounding archegonia, antheridia and
paraphyses, x 30. T. ruralis (24-32): 24. habit, x 1 ; 25. base of plant showing attachment of rhizoid gemmae,
x 3; 26. leaf (ventral surface), x20; 27. leaf with smooth costa (side view), x20; 28. leaf with scabrous costa
(side view), x20; 29. leaf in cross section (scabrous costa form), xl70; 30. upper laminal cells, x640 ; 31.
archegonia and paraphyses at stem apex, x 30; 32. rhizoid gemmae, x 170. (1-8, Magill 3820; 9-16, Ellis 3092;
17-23, Ecklon s.n.; 24, 26-27 & 30-31, Magill 6120- 25 & 32, Barnard 50349; 28-29, Magill 4036).
222
POTTIACEAE
Plants very small, forming small cushions,
green above, brownish below; terricolous.
Stems erect, 3-4 mm high, branched; in
section oval, central strand present, inner
cortical cells large, thin-walled, outer cortical
cells in 2-3 rows, smaller, incrassate, reddish.
Leaves appressed, weakly twisted around stem
dry, widespreading to recurved wet; Ungulate
to broadly ovate-lanceolate, 0, 5-1,0 mm
long; apex rounded to obtuse; margins entire,
recurved from base to near apex or infre-
quently reflexed to plane in some leaves. Costa
ending well below apex; ventral and dorsal
superficial cells rounded-quadrate above,
rectangular below, smooth; in section round,
guide cells 2-3, incrassate, slightly larger than
other cells, ventral surface cells incrassate,
smooth, dorsal substereid band of 2-3 cells,
dorsal surface cells incrassate, smooth. Upper
laminal cells rounded-quadrate to angular,
incrassate, smooth; basal cells not differen-
tiated or rectangular, thin-walled, smooth.
Gemmae infrequent, axillary, multicellular,
obovoid, 60-65 pm, brownish.
Map 85. — • Husnotiella latifolia
x Husnotiella plicata
Dioicous? Perichaetia terminal, leaves
oblong, 0, 5-1,0 mm long; apex rounded;
chlorophyllous cells in distal j of leaf. Seta
5-6 mm long, reddish yellow; capsule short-
cylindrical, 1 mm long, reddish yellow;
peristome absent; operculum rostrate, to 0,4
mm long, ± oblique; calyptra not seen;
spores round, 1 6—1 8 pm, granulate. Fig. 64:
1-9.
This species is known from Zimbabwe and South
Africa. Apparently rare, H. latifolia has been collected
in grasslands of Natal, and a few sites in the central
Cape, Orange Free State and South West Africa/
Namibia. Map 85.
Vouchers: Rehmann 459; Vahrmeijer PRE-CH
12669a.
Husnotiella latifolia is similar to the American
species H. obtusifolia (Broth.) Zander (Zander, pers.
comm.). The smooth leaf cells and non-plicate
capsules help to separate H. latifolia from the other
Southern African species.
2. Husnotiella plicata Magill, sp. nov.,
habitu H. latifolia ( Dix .) Zander & Magill
similis, sed foliis majoribus cellulis laminae
papillosis, foliis perichaetii cellulis chlorophyl-
losis restrictis ad partem distalem folii et
capsulis plicis longitudinalibus distinctis forma-
tis seriebus cellulorum dissimilium exothecii
differt.
Type: Lesotho, 210 km E. of Maseru
along Mountain Road, 1 km E. of Taung, on
soil in small canyon along Orange River,
Magill 4216 (PRE, holo.; H; MO; NY).
Plants small, in loose cushions or tufts,
green above, brownish below; terricolous.
Stems erect, 3-4 mm high, occasionally
branched; in section round, central strand
present, inner cortical cells large, thin-walled,
in 2-3 rows, outer cortical cells in 3-4 rows,
smaller, incrassate. Leaves appressed dry,
erect-spreading wet ; lingulate to oval-lanceo-
late, 0,5-1 ,2 mm long; apex acute to obtuse,
rarely rounded in some leaves; margins
generally reflexed from base to near apex,
minutely crenulate by marginal cell papillae.
Costa ending below apex; ventral superficial
cells rectangular to short-rectangular, sparsely
papillose, dorsal superficial cells rectangular,
papillose; in section round, cells scarcely
differentiated, guide cells 2, ventral cells 1-2,
in single layer, incrassate, papillose, dorsal
substereid band small, of 3-4 cells, frequently
absent, dorsal surface cells ± 6, incrassate,
papillose. Upper laminal cells rounded-hexa-
gonal, incrassate, with a single low, blunt
papilla over lumen; basal cells rectangular,
± incrassate, smooth.
Dioicous. Perichaetia quickly lateral
through innovation, leaves oblong, 1,0-1, 2
mm long; apex obtuse; chlorophyllous cells
restricted to upper ^ of leaf; lower cells
POTTIACEAE
223
Fig. 64. — Husnotiella latifolia (1-9): 1. habit,
xl; 2. habit, xlO; 3. stem in cross section, x300;
4. leaves, x40; 5. leaf in cross section, x435; 6.
cells at leaf base with attached axillary gemma (left
side), xl70; 7. upper laminal cells (dorsal surface),
x435; 8. leaf apex, Xl70; 9. perichaetial leaf, x40.
H. plicata (10—1 8) : 10. habit, xl; 11. habit, xlO;
12. stem in cross section, x300; 13. leaf, x40; 14.
leaf in cross section, x640; 15. cells at leaf base (left
side), Xl70; 16. upper laminal cells, x640; 17. leaf
apex, xl70; 18. exothecial cells at plication, x640.
(1-9, Wager PRE-CH 185; 10-18, Magill 4216).
224
POTTIACEAE
narrowly rectangular. Seta 6-9 mm long,
reddish yellow; capsule ovate to cylindrical,
narrowing to mouth, 0,8-1, 5 mm long;
exothecial cells quadrate, distinct longitudinal
plications of urn formed by linear cells in
single, vertical row; peristome absent; oper-
culum rostrate, 0,4-0, 6 mm long; calyptra
cucullate, only observed on very young
capsules; spores round, 15 p m, ± smooth.
Fig. 64: 10-18.
This new species was collected on soil in a small
kloof in semi-arid subalpine grassland above the
Orange River in central Lesotho. A subsequent
collection was made at Sehlabathebe National Park
in southeastern Lesotho. Map 85.
Vouchers: Magill 4209, 4326.
Similar to H. latifolia, this species differs in its
larger leaves with papillose leaf cells, its perichaetial
leaves with chlorophyllous cells restricted to the distal
i of the leaf, and a capsule with distinct longitudinal
plications formed by rows of differentiated exothecial
cells.
10. WEISIOPSIS
Weisiopsis Broth, in Ofvers. finska VetenskSoc. Forh. 62A: 7 (1921); Broth, in Natiirl.
PflFam. 10: 270 (1924); Sim, Bryo. S. Afr. 224 (1926); Saito in J. Hattori bot. Lab. 39: 525
(1975). Type species: W. anomala (Broth. & Par.) Broth.
Plants small, caespitose or gregarious, olive-green to brownish green; saxicolous. Stems
2-4 mm high, generally simple; central strand very small or absent. Leaves crisped to spirally
twisted with involute lamina dry, erect-spreading wet; obovate, spathulate or narrowly
elliptical, 1-4 mm long; apex rounded or acute; margins plane or involute, entire. Costa
ending below apex or percurrent; with dorsal stereid band only. Laminal cells ± rounded,
quadrate to hexagonal, incrassate, mammillose ventrally; basal cells lax. Gemmae infrequent,
cylindrical.
Monoicous or dioicous. Perichaetial leaves undifferentiated. Seta erect, 3, 5-6,0 mm long;
capsule cylindrical to ovoid, 1 ,0-1,8 mm long; peristome teeth irregularly cleft or perforated,
papillose; operculum obliquely rostrate; calyptra cucullate; spores round, granulate.
The genus Weisiopsis comprises 10 species that are found in Central America and northern South
America, the West Indies, Africa south of the Sahara, India, Asia and Oceania. The African plants grow on
rock or coarse soils from Tanzania south to Zimbabwe, Botswana and South Africa.
1 Leaf margins involute 3. W. involuta
1 Leaf margins plane:
2 Leaves broadly spathulate 1. W. plicata
2 Leaves narrowly elliptical 2. W. pulchriretis
1. Weisiopsis plicata {Mitt.) Broth, in
Ofvers. finska VetenskSoc. Forh. 62A: 8
(1921); Broth, in Natiirl. PflFam. 10: 270
(1924); Sim, Bryo. S. Afr. 224 (1926). Type:
Tanzania, Usagara Mountains, Hannington
s.n. (NY, holo. !).
Hyophila plicata Mitt, in J. Linn. Soc., Bot. 22:
304 (1886).
Plants small, gregarious or forming
small groups, olive-green, glossy; saxicolous
or terricolous. Stems 3-4 mm tall, simple; in
section round, central strand present, small,
cortical cells small, thin-walled, becoming
incrassate toward margin. Leaves crisped,
lamina widely involute dry, erect-spreading
wet; obovate, spathulate or elliptical, 1 ,5-2,5
mm long; apex obtuse to rounded; margins
plane, entire. Costa ending below apex or
percurrent; ventral superficial cells rounded-
quadrate, somewhat thickened, dorsal super-
ficial cells rectangular, incrassate; in section
round, guide cells 4, small, ventral cells in
single layer, similar to laminal cells, dorsal
stereid band 4-5 cells thick, dorsal surface
cells substereids, sometimes as large as guide
cells. Upper laminal cells rounded-quadrate,
rarely angular, incrassate, mammillose ven-
trally, smooth dorsally; basal cells large, lax,
rectangular; basal marginal cells much
smaller, quadrate to rectangular.
POTTIACEAE
225
Autoicous. Perichaetial leaves undiffe-
rentiated. Seta 3,5 mm long, yellowish;
capsule weakly plicate dry, cylindrical-ovoid,
1,6-1, 8 mm long, red-brown; peristome
irregular, teeth linear, 0,2 mm high, some-
times perforated below, granulate, light
orange; operculum rostrate, 0,5 mm long;
spores round, 10-12 pm, granulate. Fig. 65:
1-10.
Previously known only from eastern Africa,
Zimbabwe and the East African Islands, W. plicata
has recently been discovered in forests of the northern
and eastern Transvaal. Map 86.
Map 86. — • Weisiopsis plicata
x Weisiopsis pulchriretis
A Weisiopsis involuta
Vouchers: Magill 3149, 3773, 3782.
The large, spathulate leaves and distinctly plicate
capsules distinguish W. plicata from the other
Southern African species. The species could be mis-
taken for a Hyophila when sterile, but costal anatomy
will separate the two genera.
2. Weisiopsis pulchriretis Dix. in Trans.
R. Soc. S. Afr. 18: 252 (1930). Type:
Natal, Natal National Park, Wager 739 (BM,
holo. ; PRE!).
Plants small, caespitose, dark green;
saxicolous. Stems 2-4 mm long, simple; in
section without distinct central strand, corti-
cal cells small, somewhat thickened toward
margin. Leaves incurled, crisped dry, wide-
spreading wet; narrowly elliptical, 2, 5-4,0
mm long; apex acute; margins plane, entire.
Costa percurrent to mucronate; ventral
superficial cells quadrate to short-rectangular,
smooth, thin-walled, dorsal superficial cells
elongate, incrassate, smooth; in section
round, guide cells 4, ventral cells in 2-3
layers, similar to guide cells or slightly larger,
dorsal stereid band 3-4 cells thick. Upper
laminal cells hexagonal to quadrate, incras-
sate, mammillose ventrally, smooth to weakly
mammillose dorsally; basal cells rectangular
to elongate-hexagonal, thin-walled, hyaline,
narrower at margins, extending up margin to
form distinct V-shaped area.
Paroicous. Antheridia in axils of peri-
chaetial and subperichaetial leaves. Perichaetia
terminal; leaves undifferentiated. Seta 3-5
mm long, yellowish; capsule ovoid to short-
cylindrical, 1 ,0-1 ,5 mm long, yellow-brown;
peristome fragile, described as rudimentary,
only basal fragments observed; operculum
and calyptra not seen; spores round, 10-12
pm, granulate. Fig. 65: 1 1—17.
Endemic to Southern Africa, the species has
only been collected in the Goodoo Forest of Natal.
Map 86.
Voucher: Type only.
The narrowly elliptical leaves, extension of basal
leaf cells up the margins and paroicous condition
will help to identify this species. These characters
make W. pulchriretis unique in the genus. However,
the mammillose leaf cells, costal anatomy and sporo-
phyte indicate that the species is correctly placed in
Weisiopsis.
3. Weisiopsis involuta Magill, sp. nov., W.
plicata (Mitt.) Broth, habitu, anatomia costae
et cellulis laminae mammillosis ventraliter
similis, sed marginibus folii late involutis,
gemmis echinatis, setis longioribus et capsulis
non plicatis differt.
Type: Botswana, Boteti River, on near-
vertical cliff face along river, Smith 2603a
(PRE, holo.; H; MO; NY; SRGH).
Plants small, forming sods, brownish
green; saxicolous. Stems 2, 5-3, 5 mm high,
rarely branched; in section round, central
strand present, very small, inner cortical cells
large, outer cortical cells smaller, incrassate.
Leaves spirally twisted around stem to crisped,
lamina widely involute dry, erect-spreading
wet; spathulate, 0,8-1, 2 mm long; apex
ruonded; margins involute above. Costa
percurrent to ending below apex; ventral
superficial cells quadrate, thin-walled, mam-
millose, dorsal superficial cells rectangular,
incrassate, weakly prorate; in section round,
guide cells 2, ventral cells in single row,
226
POTTIACEAE
m~2.4
J-C-
'Xjrrr-f
POTTIACEAE
227
similar to laminal cells, dorsal stereid band
2-3 cells thick, dorsal surface cells substereids.
Upper laminal cells quadrate to rounded-
quadrate, incrassate, mammillose ventrally,
some cells with single, small papilla on dorsal
surface; marginal cells frequently with several
papillae; basal cells larger, quadrate to
rectangular, smooth. Gemmae infrequent,
cylindrical, echinate, 150-200 pm long.
Dioicous. Plants undifferentiated. Peri-
gonium terminal, perigonial leaves with oval
bases. Perichaetia terminal, perichaetial leaves
lingulate, 1 ,0-1,5 mm long. Seta 4, 5-6, 5
mm long, brownish; capsule ovate to ellip-
tical, 1,0-1, 2 mm long, brownish yellow;
peristome yellowish, teeth irregularly cleft
above, fragile, united at base forming a short
basal membrane, papillose; operculum obli-
quely rostrate, 0,5 mm long; spores round,
10-12 pm, granulate. Fig. 65: 18-27.
This new species is presently known only from
northern Botswana. The type locality is a near-vertical,
soft, crumbly calcrete cliff along the Boteti River.
Map 86.
Voucher: Type only.
The small size of the plants, broadly involute
leaf margins, production of gemmae, long seta, and
smooth capsules separate W. involuta from other
species of Weisiopsis. The spathulate leaves, costal
anatomy, small mammillose laminal cells and presence
of a peristome indicate that the species belongs in
Weisiopsis.
Tribe BARBULEAE
Plants small to medium, in loose tufts; terricolous or saxicolous. Stems erect; centra
strand present. Leaves frequently board; lanceolate to ovate-ligulate or elliptical to oblong;
margins plane to recurved or revolute. Costa ending below apex to long-excurrent; generally
with dorsal and ventral stereid bands. Laminal cells generally thickened, smooth to mammillose
or papillose.
Capsule stegocarpic; peristome present or absent, teeth generally filiform, erect or twisted;
operculum conic to rostrate; calyptra cucullate.
Key to Genera of Tribe Barbuleae
1 Leaves broadly elliptical ; apex broadly acute to obtuse ; margins narrowly involute when
dry; laminal cells strongly mammillose ventrally; peristome absent 1. Hyophila
1 Leaves narrower, margins plane to revolute wet or dry; laminal cells generally papillose
or mammillose on both dorsal and ventral surfaces:
2 Upper leaf margins bistratose 2. Trichostomopsis
2 Upper leaf margins unistratose:
3 Plants generally reddish green; leaf apices with a distinct, clear, smooth apiculus
and occasionally one to several smooth, clear marginal teeth; basal leaf cells
strongly differentiated 5. Bryoerythrophyllum
Fig. 65. — Weisiopsis plicata (1-10): 1. habit, x 1; 2. habit, x 10; 3. stem in cross section, x 150; 4. leaves,
x20; 5. leaf in cross section, x225; 6. basal leaf cells (right side), xl70; 7. upper laminal cells, xl70; 8. leaf
apex, xl70; 9. perichaetial leaf, x20; 10. part of capsule mouth with peristome teeth and spores, xl70.
W. pulchriretis (11-17): 11. habit, xl; 12. habit, x 10; 13. stem in cross section, xl50; 14. leaves, x20; 15.
leaf in cross section, x225; 16. cells at leaf base (left side), x 170; 17. leaf apex, x 170. W. involuta (18-27):
18. habit, x 1; 19. habit, x 10; 20. stem in cross section, x200; 21. leaf, x40; 22. leaf in cross section, x260;
23. cells at leaf base (right side), x435; 24. leaf apex, x435; 25. upper laminal cells, x 640; 26. axillary gemmae,
x220; 27. part of capsule mouth with peristome teeth, xl70. (1-10, Magill 3782; 1 1—17, Wager 739; 18-27,
Smith 2603a).
228
POTTIACEAE
3 Plants green to brownish green; leaf apices without smooth, clear apiculus or margi-
nal teeth; basal cells usually somewhat differentiated:
4 Axillary hairs with basal cell brownish; ventral superficial costal cells quadrate
to short-rectangular 3. Didymodon
4 Axillary hairs hyaline throughout; ventral superficial costal cells rectangular
to elongate 4. Barbula
1. HYOPHILA
Hyophila Brid., Bryo. Univ. 1: 760 (1827); Broth, in Natiirl. PflFam. 10: 269 (1924);
Gangulee, Moss. E. India 677 (1972). Lectotype species: H. javanicum (Nees & Blum.)
Brid., vide Hampe in Bot. Ztg 4: 266 (1846).
Plants small to medium, caespitose; saxicolous or terricolous. Stems erect, little branched;
central strand present. Leaves glossy, incurled with involute margins dry, spreading with plane
margins wet; broadly oblong or elliptical, sometimes spathulate; apex broadly acute to
rounded, mucronate; margins plane, entire or serrate to dentate. Costa percurrent or short-
excurrent; in section with dorsal and ventral stereid bands. Laminal cells small, quadrate or
hexagonal, mammillose ventrally, smooth dorsally; basal cells short-rectangular to quadrate,
smooth. Gemmae infrequent, axillary, polymorphous, multicellular.
Dioicous. Perichaetial leaves undifferentiated. Seta short, 8-12 mm long; capsule cylin-
drical, gymnostomous; operculum rostrate; calyptra cucullate; spores round, essentially
smooth.
The approximately 110 species of Hyophila are most common in the tropics. A few, rather widespread
species, e.g. H. involuta, are also present in temperate regions. In Southern Africa the genus is found in Natal
and a few locations in the eastern Transvaal.
1 Leaves 2,4 mm long, margins serrate to dentate 1. H. involuta
1 Leaves 1,0-1, 5 mm long, margins entire or rarely serrulate 2. H. baginsensis
1. Hyophila involuta (Hook.) Jaeg. in
Verh. St Gall, naturw. Ges. 1 871—72 : 354
(1873); Saito in J. Hattori bot. Lab. 39: 468
(1975); Gangulee, Moss. E. India 681 (1972).
Type: Nepal, D. Gardner s.n.
Gymnostomum involutum Hook., Muse. Exot. 2:
154 (1819).
Trichostomum atrovirens Rehm. ex C. Mull, in
Hedwigia 38: 100 (1899). Hyophilia atrovirens (C.
Mull.) Broth, in Natiirl. PflFam. 1: 403 (1902);
Sim, Bryo. S. Afr. 222 (1926). Type: Natal, Van
Reenens Pass, Rehmann 119 (PRE!).
Plants small to medium, in loose tufts,
blackish green to dark green; saxicolous.
Stems 5-15 mm tall; in section round, central
strand present, inner cortical cells thin-walled,
becoming smaller toward margin, incrassate,
reddish. Leaves glossy, incurled with involute
margins dry, spreading with plane margins
wet; broadly oblong to elliptical, 2-4 mm
long; apex broadly acute, mucronate; margins
plane, serrate to dentate in upper leaf. Costa
subpercurrent to short-excurrent, ventral
superficial cells quadrate, mammillose, dorsal
superficial cells rectangular, smooth, incras-
sate; in section round to oval, guide cells 4,
small, ventral stereid band small, 2-3 cells
thick, ventral surface cells thickened, bulging,
dorsal stereid band large, 2-6 cells thick,
dorsal surface cells substereids. Upper laminal
cells small, hexagonal to subhexagonal or
rounded, becoming quadrate below; in sec-
tion ventral surface mammillose, dorsal
surface smooth; basal cells rectangular,
smooth. Gemmae infrequently produced,
axillary, stalked, multicellular, polymorphous.
Dioicous. Perichaetia terminal, leaves
undifferentiated. Seta erect, to 8 mm long,
yellowish; capsule cylindrical, 1,5-1, 8 mm
long, brownish; peristome absent; operculum
rostrate, 0,8 mm long; calyptra cucullate;
spores round, 10-13 pm, smooth. Fig. 66:
1-9.
POTTIACEAE
229
Fig. 66. — Hyophila involuta (1-9): 1. habit, wet,
X 1 ; 2. habit, dry, x 1 ; 3. habit, x 10; 4. stem in cross
section, x 170; 5. leaves, X 20; 6. leaf in cross section,
x200; 7. basal leaf cells (right side), x 170; 8. upper
laminal cells, x640 ; 9. leaf apex, xl70. H. bagin-
sensis (10-16): 10. habit, xl; 11. habit, xlO; 12.
leaves, x20; 13. leaf in cross section, x200; 14.
upper laminal cells, x640; 15. leaf apex, xl70; 16.
axillary gemmae, x200. (1-9, Sim PRE-CH5816;
10-16, Magill 4924).
230
POTTIACEAE
New to Africa, Hyophila involuta was previously
known from Europe, southern Asia and India,
Micronesia and North, Central and South America.
It is probable that the species occurs elsewhere in
Africa and re-examination of tropical African speci-
mens, especially central African collections of H.
atrovirens, may provide additional records for this
species. In Southern Africa the species is found in
forests or grasslands of the Orange Free State,
Lesotho, Natal and eastern Transvaal. Map 87.
Map 87. — • Hyophila involuta
x Hyophila baginsensis
Vouchers: Crosby & Crosby 7866; Rankin 199a;
Scheepers 1217.
The broad, oblong leaves and serrate to dentate
margins will help to identify this species. The other
Southern African species has spathulate leaves with
more or less entire margins.
2. Hyophila baginsensis C. Mull, in
Linnaea 39: 399 (1875); Broth, in Natiirl.
PflFam. 10: 270 (1924); Sim, Bryo. S. Afr.
222 (1926). Type: Central Africa, Niam Niam
Region, Bagins s.n., 27 May 1870.
Plants small, caespitose, dark green to
yellowish green; saxicolous or terricolous.
Stems to 6 mm tall; in section round, central
strand present, cortical cells thickened, outer
row smaller, incrassate, reddish. Leaves
glossy, incurled with involute margins dry,
widespreading with plane margins wet;
broadly elliptical to lingulate or spathulate,
1,0-1, 5 mm long; apex rounded to obtuse,
mucronate; margins plane, entire or rarely
serrulate above. Costa very short-excurrent,
ventral superficial cells quadrate to short-
rectangular, mammillose, dorsal superficial
cells linear, incrassate, smooth; in section
round, guide cells 4, ventral stereid band 1-2
cells thick, ventral surface cells larger,
incrassate, dorsal stereid band large, 4-6 cells
thick, dorsal surface cells substereids. Upper
laminal cells small, rounded-quadrate, mam-
millose ventrally, smooth dorsally; basal cells
short-rectangular to quadrate, thin-walled.
Gemmae axillary, stalked, stellate, multicel-
lular.
Sporophyte not known from Southern
Africa. Described from Zimbabwe (Sim,
1926) as: ‘Seta to 12 mm long; capsule
cylindrical, contracted at mouth; peristome
absent’. Fig. 66: 10-16.
Growing on soil, H. baginsensis occurs in shrub-
savanna and forests of central, eastern and Southern
Africa. Within the Flora area the species is found in
Zululand, Swaziland, and the northern and eastern
Transvaal. Map 87.
Vouchers: Magill 4746, 4924; Sim 7512.
The smaller size and generally entire leaf margins
of H. baginsensis distinguish it from H. involuta. The
species could be confused with Weisiopsis but
presence of a ventral stereid band in the costa and
absence of a peristome characterize the species.
2. TRICHOSTOMOPSIS
Trichostomopsis Card, in Rev. Bryol. 36: 73 (1909); Broth, in Natiirl. PflFam. 10: 264 (1924);
Robinson in Phytologia 20: 184 (1970). Type species: T. crispifolia Card.
Barbula section Asteriscium C. Miill. in Linnaea 42: 342 (1872); Sim, Bryo. S. Afr. 238 (1926).
Plants small, caespitose; terricolous. Stems erect, to 10 mm tall, simple; central strand
present. Leaves lanceolate to ovate; apex acute; margins plane, entire, bistratose. Costa
percurrent; in section with dorsal stereid band only. Laminal cells quadrate to short-rectan-
gular, incrassate, smooth or pleuropapillose; marginal cells smaller; basal cells rectangular,
thin-walled; basal marginal cells narrowly rectangular.
POTTIACEAE
231
Fig. 67. — Trichostomopsis australasiae (1-9): 1.
habit, xl; 2. habit, x 10; 3. stem in cross section,
X 130; 4. leaves, x40; 5. leaf in cross section, x435;
6. cells at leaf base (left side), x 170; 7. upper laminal
cells at left margin, xl70; 8. leaf apex, xl70; 9.
part of capsule mouth with peristome teeth, xl70.
T. trivialis (10-16): 10. habit, xl; 11. habit, xlO;
12. stem in cross section, xl30; 13. leaves, x40; 14.
leaf in cross section, x435; 15. cells at leaf base
(right side), x 170; 16. leaf apex, x 170. (1-9, Schelpe
7803; 10—16, Rehmann 99).
232
POTTIACEAE
Dioicous. Capsule short, cylindrical; peristome teeth divided into 2 linear, papillose
filaments, yellowish; operculum rostrate; calyptra cucullate, smooth; spores subround,
smooth to granulate.
Trichostomopsis australasiae is a widespread species known from drier parts of both hemispheres. The
other four species of Trichostomopsis are. very restricted in distribution. In Southern Africa, the genus is found in
semi-arid regions of the Cape and Orange Free State.
The genus was recently put into the synonomy of Didymodon by Zander (1978).
1 Leaf cells papillose
1 Leaf cells smooth
1. Trichostomopsis australasiae (Hook. &
Grev.) Robinson in Phytologia 20: 187 (1970).
Type: Australia, King George’s Sound,
Menzies s.n., 1791.
Tortula australasiae Hook. & Grev. in Edinb. J.
Sci. 1: 301 (1824). Barbula australasiae (Hook. &
Grev.) Brid., Bryol. Univ. 1: 828 (1827); Broth, in
Natiirl. PflFam. 10: 278 (1924).
Barbula trichostomacea C. Miill. in Hedwigia 38:
108 (1899); Sim, Bryo. S. Afr. 238 (1926). Type: Cape,
Rondebosch, Rehmann 97 (PRE!).
Plants small, caespitose, dark green;
terricolous. Stems to 10 mm high, simple; in
section round, central strand small, inner
cortical cells large, weakly thickened, outer
cortical cells in 1-2 rows, smaller, incrassate,
reddish. Leaves incurved to contorted dry,
spreading wet; lanceolate, 1-2 mm long,
lamina unistratose, rarely with small bistra-
tose areas or bistratose apically; apex acute;
margins bistratose above base. Costa per-
current, ventral superficial cells quadrate,
papillose, dorsal superficial cells short- to
long-rectangular, smooth; in section oval,
guide cells 2-4, ventral cells in l(-2) layer(s),
incrassate, papillose, dorsal stereid band 1-2
cells thick, dorsal surface cells substereids.
Upper laminal cells subquadrate, rounded,
with 1-4 low, blunt papillae per cell, incon-
spicuous; basal cells rectangular, thin-walled;
basal marginal cells frequently narrower.
Dioicous. Perichaetial leaves undifferen-
tiated. Seta 7-8 mm long, dark red; cap-
sule short-cylindrical, 1, 5-2,0 mm long;
peristome teeth divided to near base, filaments
erect, finely papillose; operculum conic, 0,6
mm long; spores subround, 12-17 pm,
smooth. Fig. 67: 1-9.
Trichostomopsis australasiae occurs in North,
Central and South America, Southern Africa,
Australia and New Zealand. In Southern Africa the
species is found in the western Cape, Orange Free
State, Lesotho, eastern Transvaal and central South
West Africa/Namibia. Map 88.
1. T. australasiae
. . . 2. T. trivialis
Map 88. — • Trichostomopsis australasiae
X Trichostomopsis trivialis
Vouchers: Garside 6505; Magill & Schelpe 3880d;
Phelan et al. 726; Schelpe 7755.
The presence of minute leaf cell papillae, although
frequently only obvious in transverse sections of the
leaves, help to separate this species from T. trivialis.
The bistratose margins distinguish T. australasiae from
other related species in the tribe.
2. Trichostomopsis trivialis (C. Mull.)
Robinson in Phytologia 20: 187 (1970). Type:
Orange Free State, Kadziberg, Rehmann 99
(PRE!).
Barbula trivialis C. Mull, in Hedwigia 38: 107
(1899).
Plants small, caespitose, green to yellow-
green; terricolous. Stems to 10 mm high,
simple; in section round, central strand very
small, inner cortical cells lax, outer cortical
cells in 1-2 rows, small, incrassate, reddish.
Leaves incurled dry, erect wet; lanceolate to
ovate-acuminate, 1, 5-2,0 mm long, lamina
unistratose; apex broadly acute; margins
bistratose, cells slightly smaller than laminal
cells. Costa percurrent, ventral superficial
POTTIACEAE
233
cells quadrate, smooth, dorsal superficial cells
rectangular, smooth; in section oval, guide
cells 4-5, ventral cells in l(-2) layer(s), large,
mammillose, dorsal stereid band 2-3 cells
thick, dorsal surface cells undifferentiated.
Upper laminal cells irregularly quadrate to
short-rectangular, smooth or nearly so ;
marginal cells quadrate, smaller; basal cells
rectangular, thin-walled ; basal marginal cells
in 1-3 rows, narrowly rectangular.
Dioicous. Perichaetial leaves undifferen-
tiated. Seta 8-10 mm long, reddish yellow;
capsule short-cylindrical, 1,5 mm long;
peristome divided to base, filaments erect,
papillose, yellowish red; operculum rostrate,
0,8 mm long; spores subround, 15 pm,
granulate. Fig. 67: 10-16.
Endemic to Southern Africa, the species is known
from the central Cape, Orange Free State and Lesotho.
The label on the type specimen gives Kadziberg,
Orange Free State. Re-examination of Rehmann’s
route indicates that the Kadziberg is part of the
Witteberg in the northeastern Orange Free State, near
the Caledon River. Map 88.
Voucher: Magill 4255.
The species differs from T. australasiae in leaf
shape, anatomy and smooth leaf cells.
3. DIDYMODON
Didymodon Hedw., Spec. Muse. 104 (1801); Saito in J. Hattori bot. Lab. 39: 500 (1975).
Lectotype species: D. rigidulus Hedw., vide Grout, Moss FI. N. Amer. 1: 186 (1939).
Plants small to medium, forming cushions, yellowish green to dark green, reddish or
brownish below; terricolous or saxicolous. Stems with central strand; axillary hairs short,
basal cells distinct, brownish. Leaves appressed, contorted or spirally twisted dry, spreading
wet; lanceolate to ligulate; apex acute to rounded; margins entire, reflexed to revolute.
Costa ending below apex or short-excurrent; in section ventral cells incrassate to substereids,
dorsal stereid band strong. Laminal cells rounded-quadrate, mammillose to weakly papillose;
basal cells rectangular, smooth.
Dioicous. Perichaetia terminal. Seta 7-20 mm long; capsule cylindrical; peristome
present or absent; operculum rostrate; calyptra cucullate.
The genus Didymodon comprises c. 100 species and is found in temperate and tropical regions of continents
in both hemispheres (one species reported from Antarctica). The major concentration of species is found in
Central and South America.
Saito (1975) considered Didymodon a natural genus, closely related to Barbula. Another related genus,
Trichostomopsis, was recently considered to be a synonym of Didymodon by Zander (1978). Although the
separation of the two genera is not always clear-cut, Trichostomopsis has been maintained here. Didymodon is
separated from Barbula by differentiated basal cells of the axillary hairs, quadrate to short-rectangular
ventral superficial costal cells and weakly papillose leaf cells. The reflexed to revolute, unistratose leaf margins
separate Didymodon from Trichostomopsis.
1 Leaf apices acute to rounded on same plant ; margins plane above, reflexed below 1 . D. ceratodonteus
1 Leaf apices acute to acuminate throughout; margins revolute to spirally revolute in upper leaf
2. D. xanthocarpus
1. Didymodon ceratodonteus (C. Mull.)
Dix. in K. norske Vidensk. Selsk. Skr.
1932 (4): 7 (1932). Type: Cape, Philipstown
at Kat River, Ecklon s.n. (BM!).
Pottia ceratodontea C. Mull., Syn. Muse. 1 : 564
(1849). Hymenostylium ceratodonteum (C. Mull.)
Broth, in Natiirl. PflFam. 1: 389 (1902); Sim, Bryo.
S. Afr. 256 (1926).
Trichostomum afrum C. Mull, in Hedwigia 38: 98
(1899). Didymodon afer (C. Mull.) Broth, in Natiirl.
PflFam. 1: 406 (1902). Type: Cape, Somerset East,
Boschberg, MacOwan s.n. (Herb. Winter).
Barbula dimorpha C. Miill. in Hedwigia 38: 106
(1899). Didymodon dimorphum (C. Miill.) Broth, in
Natiirl. PflFam. 1 : 407 (1902). Gymnostomum dimor-
phum (C. Mull.) Sim, Bryo. S. Afr. 259 (1926). Type:
Cape, Cape Town, Rehmann 102 (BM!).
Didymodon pottsii Dix. in Bull. Torrey bot. Club
43: 193 (1916); Gymnostomum pottsii (Dix.) Sim,
Bryo. S. Afr. 260 (1926). Type: Orange Free State,
Eagle’s Nest, Bloemfontein, Potts s.n. (Sim 8663;
PRE!).
Weissia gracilis Wag. & Dix. in Trans. R. Soc.
S. Afr. 4: 4 (1914). Gymnostomum gracile (Wag. &
Dix.) Dix. in Trans. R. Soc. S. Afr. 8: 191 (1920),
234
POTTIACEAE
Fig. 68. — Didymodon xanthocarpus (1-10): 1.
habit, xl; 2. habit, xlO; 3. stem in cross section,
x 170; 4. leaves, x40; 5. leaf in cross section, x435;
6. cells at upper leaf base (right side), x 170; 7. upper
laminal cells, x640; 8. leaf apex, X 170; 9. perichae-
tial leaf, x40; 10. peristome teeth and capsule mouth,
x40. D. ceratodonteus (11-19): 11. habit, xl; 12.
habit, xlO; 13. stem in cross section, x220; 14-15.
leaves, x40; 16. leaf in cross section, x435; 17.
cells at mid-leaf (right side), X 170; 18. upper laminal
cells, x640; 19. leaf apex, Xl70. (1-2, Magill 6241;
3-10, Magill 3856; 11-19, Relief 366).
POTTIACEAE
235
non G. gracile (R. Br.) Hook., Musci Exot. 1 : 22
(1818). Gymnostomum wageri Schelpe in Mem. bot.
Surv. S. Afr. 43: 5 (1979). Type: Transvaal, Pretoria,
Wager 97 (PRE!).
Hyophila basurensis Sim, Bryo. S. Afr. 223 (1926).
Type: Cape, Renosterberg, Rehmann 458 (PRE,
holo.!).
Didymodon perrevolutus P. Varde in Revue bryol.
lichen. 27: 4 (1958). Type: Cape, Partridge Point,
Cape Peninsula, Arnell 1188 (PC, holo.!).
Plants small to medium, in dense
cushions, light green above, light brown
below; terricolous or saxicolous. Stems 10-50
mm tall, frequently branched, especially in
taller plants; in section round, central strand
small, inner cortical cells large, thin-walled,
outer cortical cells in 2-3 rows, smaller,
incrassate, yellowish, outermost row sub-
stereids. Leaves appressed to slightly con-
torted dry, patent wet, shape extremely
variable on same plant, ovate-acuminate to
lanceolate or ligulate, 1,0-1, 8 mm long;
apex acute to broadly rounded; margins
entire, reflexed to recurved below, plane to
reflexed above. Costa percurrent to ending
10 cells below rounded apices, ventral and
dorsal superficial cells quadrate to short-
rectangular, incrassate, smooth; in section
round, weakly differentiated, guide cells 2,
ventral cells in 1-2 layers, incrassate, dorsal
stereid band 1-2 rows thick, frequently cells
substereids only, dorsal surface cells larger,
similar to ventral cells, incrassate. Upper
laminal cells quadrate to rounded-quadrate,
incrassate, frequently with thickened corners
in upper leaf, some cells with low, blunt
papillae frequently obvious only in leaf
sections; basal cells rectangular to short-
rectangular, thin-walled, smooth.
Perichaetia terminal, becoming lateral
through innovation, leaves ligulate, 1,5 mm
long; apex obtuse to broadly rounded; leaf
cells occasionally somewhat larger than
vegetative leaf cells. Seta erect, 7-8 mm long,
reddish; capsule short-cylindrical, 1,5 mm
long, reddish to yellowish red; peristome
absent; operculum rostrate, 0,4 mm long;
spores round, 12-15 pm, weakly granulate.
Fig. 68: 1 1-19.
Didymodon ceratodonteus is presently known
from Zimbabwe, South West Africa/Namibia, Bo-
tswana, the central, eastern and southern Transvaal,
Orange Free State, Lesotho, Zululand, Natal and the
eastern, central and southwestern Cape. Map 89.
Vouchers: Cholnoky 768; Retief 366a.
Map 89.— • Didymodon ceratodonteus
Variation in size, leaf shape and rounding of the
leaf apex account for the synonomy accompanying
this species. Although many of the plants appear
distinct macroscopically, the costal anatomy and cell
size, shape and patterns are very consistent. In
addition, a careful examination will clearly demon-
strate the variation expressed by the leaves and leaf
apices of most plants.
2. Didymodon xanthocarpus (C. Mull.)
Magill in Mem. bot. Surv. S. Afr. 43: 5
(1979). Type: Cape, Swartkop River, Ecklon
s.n. (BM!; H-SOL!).
Barbula xanthocarpa C. Mull, in Linnaea 17: 581
(1843).
Didymodon knysnae Rehm. ex Sim, Bryo. S. Afr.
248 (1926). Type: Cape, Knysna, Rehmann 83 (PRE,
holo.!).
Plants medium-sized, forming cushions,
yellowish green to reddish or dark green
above, reddish below, frequently glossy;
terricolous. Stems 10-20 mm tall, frequently
branched above; in section round, central
strand present, generally large, inner cortical
cells large, incrassate, reddish or yellowish,
outer cortical cells in 2-3 rows, stereids, red.
Leaves contorted or rarely twisted around
stem dry, widespreading wet; lanceolate to
oval-acuminate, 2, 0-3, 5 mm long, keeled,
rarely bistratose above; apex acute, mucro-
nate; margins entire, revolute throughout.
Costa strong, short-excurrent, ventral super-
ficial cells rounded-quadrate, incrassate,
smooth, dorsal superficial cells rectangular,
incrassate, smooth; in section round, semi-
circular or oval, guide cells 4-6, ventral cells
236
POTTIACEAE
in 2-3 layers, similar to guide cells or fre-
quently incrassate, ventral surface cells
incrassate, mammillose or papillose, dorsal
stereid band strong, 4-5 cells thick, reddish,
dorsal surface cells larger, outer walls more
strongly thickened. Upper laminal cells small,
rounded-quadrate, incrassate, bulging or
some cells mammillose; basal cells elongate,
thin-walled, smooth.
Dioicous. Perichaetia terminal, leaves
oblong, 2,0-2, 5 mm long; apex irregularly
notched; laminal cells irregularly shaped,
triangular to rectangular, incrassate, smooth.
Seta erect, 8-20 mm long, reddish yellow
to dark red; capsule cylindrical to oblong-
elliptical, 1,5-2, 5 mm long, constricted at
mouth; peristome weakly twisted, teeth
linear, 0,8 mm long, finely papillose, yellow-
ish red; operculum rostrate, 1,5 mm long;
calyptra cucullate; spores round, 12,5-15,0
pm, granulate to smooth. Fig. 68: 1-10.
Known only from Southern Africa, D. xantho-
carpus is found in shrublands of the Cape Province.
Map 90.
Vouchers: Claassen 255; Esterhuysen 20148;
Magill 5852; Schelpe 7807.
The lanceolate leaves with revolute margins and
mammillose leaf cells will help to identify this species.
The anatomy of the costa, with ventral cells in several
rows, large and little thickened, is also distinctive.
A few specimens with bistratose leaves have been
collected from the semi-arid regions of the western
Cape. The bistratose leaves, frequently encountered
in Pottiaceae, are seen as an adaptation to the dry
habitat. The specimens conform in all other characters
to the unistratose-leaved plants. Another modifica-
tion observed in a few collections, was the presence of
several weak papillae on the leaf cells. These plants
generally do not have the strongly thickened mammil-
lose leaf cells. It is possible that the bulging thicken-
ings may obscure the fine papillae in most specimens.
4. BARBULA
Barbula Hedw., Spec. Muse. 115 (1801); Saito in J. Hattori bot. Lab. 39: 481 (1975). Type
species: B. unguiculata Hedw.
Trichostomum section Hydrogonium C. Miill. in Linnaea 40: 297 (1876). Hydrogonium (C. Miill.) Jaeg.,
Adumbratio 2: 669 (1880). Lectotype species: H. ehrenbergii (Lor.) Jaeg., vide Saito in J. Hattori bot. Lab. 38:
492 (1975).
Semibarbula Herz. ex Hilp. in Beih. bot. Zbl. 50: 626 (1933). Type species: S. indica (Brid.) Hilp.
Plants small to medium, in loose tufts or cushions, yellow-green to dark green; terri-
colous or saxicolous, infrequently aquatic. Stems erect, flaccid if aquatic, sparsely branched;
in section round, central strand present, sometimes weak; axillary hairs 4-10 cells long, hyaline
throughout. Leaves contorted or spirally twisted dry, spreading wet, plane or rugose; lanceo-
late to ligulate or ovate-ligulate; apex acute to obtuse; margins plane to spirally revolute,
entire, generally papillose. Costa ending below apex to long-excurrent, ventral superficial
cells rectangular to elongate, dorsal superficial cells rectangular, smooth, prorate or papillose;
in section with dorsal and ventral stereid bands at least in lower leaf. Laminal cells quadrate
to hexagonal, papillose to nearly smooth; marginal cells sometimes differentiated; basal cells
weakly to strongly differentiated, elongate, smooth or papillose. Gemmae axillary or on rhi-
zoids, multicellular, obovoid.
POTTIACEAE
237
Dioicous. Perichaetia terminal; leaves larger, generally with sheathing base. Seta erect;
capsule cylindrical; peristome teeth linear above short basal membrane, twisted; operculum
conical; calyptra cucullate; spores round, smooth to papillose.
The genus Barbuta, including Semibarbula and Hydrogonium, comprises c. 300 species and is the largest
genus in the family. The genus is well represented in temperate regions of both hemispheres and is found on all
continents, including Antarctica. The major concentration of species is found in western North and South
America.
1 Costa smooth dorsally:
2 Leaf margins plane in upper leaf:
3 Leaves rugose; costa mucronate to aristate 8. B. eubryum
3 Leaves smooth; costa ending in or below apex 9. B. afrofontana
2 Leaf margins revolute to spirally revolute in upper leaf:
4 Costa long-excurrent as a smooth, yellow awn 1. B. crinita
4 Costa mucronate to aristate:
5 Leaves ligulate to lanceolate; marginal cells surrounded by spiralling lamina differentiated into
specialized photosynthetic region 2. B. acutata
5 Leaves ovate-acuminate; marginal cells undifferentiated 3. B. hornschuchiana
1 Costa scabrous or papillose dorsally:
6 Basal leaf cells strongly differentiated, long-rectangular, yellowish, seriate papillose at junction with
upper laminal cells:
7 Leaves lanceolate to ligulate, 2-3 mm long; seta to 50 mm long, operculum very long 4. B. calycina
7 Leaves ligulate above obovate base, 1 ,5-2,5 mm long; seta to 15 mm long, operculum short
5. B. microcalycina
6 Basal leaf cells short-rectangular, smooth:
8 Costa mucronate, dorsal superficial cells prorate 6. B. indica
8 Costa ending below apex, dorsal superficial cells seriate papillose 7. B. rehmannii
1. Barbula crinita Schultz in Nova Acta
Acad. Leop. Carol. 11: 226 (1823). Type:
Java, Dickson s.n.
Tortula pilifera Hook., Muse. Exot. 1: 12 (1818);
Sim, Bryo. S. Afr. 228 (1926). Barbula pilifera (Hook.)
Brid., Bryo. Univ. 1: 572 (1826), horn, illeg. ; Broth,
in Natiirl. PflFam. 10: 280 (1924); Weber in Lind-
bergia 1:214(1972).
Tortula pilifera Hook. var. longifolia Sim, Bryo. S.
Afr. 229 (1926). Type: Cape, King William’s Town,
Sim 7323 (PRE, holo. !).
Barbula torquescens Schimp. ex C. Mull, in Bot.
Ztg 16: 163 (1858); Broth, in Natiirl. PflFam. 10: 280
(1924). Type: Cape, Groenkloof, Breutel s.n. (BM!).
Plants medium-sized, in loose tufts,
light green to yellow-green; terricolous or
saxicolous. Stem to 35 mm high, infrequently
branched; in section round, central strand
small, cortical cells lax becoming incrassate
near margin, 1-3 marginal rows smaller,
substereids. Leaves contorted to spirally
twisted dry, erect- to widespreading wet;
lanceolate to linear above ovate base, 2,0-
4,5 mm long; apex broadly acute to obtuse,
frequently asymmetrical; margins recurved
from base to apex, entire. Costa long-excur-
rent as yellowish, smooth to weakly denticu-
late awn, 1-2 mm long, ventral superficial
cells quadrate, papillose, dorsal superficial
cells elongate, strongly incrassate, smooth; in
section reniform, guide cells 4, ventral
stereid band small, over central guides, 2-3
cells thick, infrequently cells substereids,
ventral surface cells thin-walled, papillose,
dorsal stereid band large, 4-6 cells thick,
dorsal surface cells unevenly thickened,
lumen crescent-shaped. Upper laminal cells
hexagonal to quadrate, with 2-4 large,
C-shaped papillae over lumen; basal leaf
cells short- to long-rectangular, smooth.
Dioicous. Perichaetial leaves sheathing
seta, oblong to elliptical, 4-5 mm long;
apex acute; margins plane, entire, laminal
cells elongate, rectangular to polygonal;
basal cells long-rectangular. Seta erect, to
20 mm long, reddish yellow; capsule cylin-
drical, to 4 mm long, reddish brown; peri-
stome yellowish, teeth filiform, papillose,
spirally twisted, basal membrane short, 1-3
238
POTTIACEAE
POTTIACEAE
239
cells above mouth; operculum conical, 1,5
mm long; spores round, 10-12 pm, finely
papillose. Fig. 69: 1-10.
Barbula crinita is a rather common moss in dry
shrublands of Southern Africa, South America,
Australia and New Zealand. The species is common
throughout semi-arid and arid regions of the Cape and
southern South West Africa/Namibia, but is also
found in the Orange Free State, Lesotho, Natal and
Transkei. A few specimens are known from the eastern
and northern Transvaal. Map 91.
Map 91. — • Barbula crinita
Vouchers: Boucher PRE-CH12771; Esterhuysen
20585; Garside 6614; Hardy 4219a; Lavranos 15554c;
Magill 4655, 5817, 6049; Schelpe 7795; Van Rooy 37.
The general habit of this species and the long-
excurrent costa, indicate a relationship to Tortula.
The presence of a ventral stereid band in the costa
clearly indicates that the species is a Barbula.
2. Barbula acutata C. Mull, in Hedwigia
38: 109 (1899); Broth, in Natiirl. PflFam.
10: 280 (1924). Type: Transvaal, Spitzkop,
near Lydenburg, Wilms s.n., 1887 (G,
holo. !).
Plants medium-sized, in loose cushions,
green; terricolous. Stems 8-15 mm high; in
section with large central strand, inner
cortical cells in 3-4 rows, lax, outer cortical
cells in 1-2 rows, smaller, incrassate, reddish.
Leaves appressed, spirally twisted around
stem dry, erect-spreading wet; broadly
lanceolate to ligulate, 1,0-1, 5 mm long;
apex obtuse to broadly acute, mucronate;
margins spirally revolute, entire, marginal
cells differentiated into specialized photo-
synthetic region; in section juxtacostal 1am-
inal cells papillose on dorsal and ventral
surfaces, cells decreasing in height toward
revolute margins; cells smooth dorsally in
revolute portion, 4-6 marginal cells enclosed
by spiralling leaf, enlarged, thin-walled.
Costa short-excurrent to percurrent, strong
to apex, 60-70 /am wide at apex, ventral
superficial cells quadrate, papillose, dorsal
superficial cells elongate-rectangular, incras-
sate, smooth; in section reniform to flattened,
guide cells 4-5, large, incrassate, ventral
stereid band small, 1-4 cells over central
guide cells, ventral surface cells larger, thin-
walled, papillose, dorsal stereid band strong,
5-6 cells thick, dorsal surface cells with
strongly thickened outer walls, lumen lunate.
Upper lamina! cells quadrate to short-
rectangular, with 4-6 low, O- or C-shaped
papillae over lumen; marginal cells differen-
tiated, enlarged, smooth; basal cells rectangu-
lar, smooth.
Sporophyte not known. Fig. 69: 11-19.
Barbula acutata is found only in Southern Africa.
The species is associated with high, semi-arid grass-
lands of central South West Africa/Namibia, Botswa-
na, the northern, eastern and central Transvaal,
Orange Free State and Lesotho. Map 92.
Vouchers: Magill 3637, 4256; Phelan et al. 72a;
Schelpe PRE-CHI 2750; Van Rooy 329.
The highly specialized marginal cells and presence
of the ventral stereid band in the costa characterize
this species. These features also place the species
within Zander’s (1979) concept of Pseudocrossidium
Williams. The presence of specialized marginal cells
in Tortula porphyreoneura suggests that modification
Fig. 69. — Barbula crinita (1-10): 1. habit, wet, x 1 ; 2. habit, dry, x 1 ; 3. habit, x 5; 4. stem in cross section,
x 200; 5. leaves, x 20; 6. leaf in cross section, x 170; 7. cells at leaf base (right side), x 170; 8. upper laminal
cells, x640; 9. leaf apex, xl70; 10. capsule mouth with peristome, x25. B. acutata (11-19): 11. habit, xl;
12. habit, x5; 13. stem in cross section, x200; 14-15. leaves, x40; 16. leaf in cross section, x300; 17. basal
leaf cells (right side), xl70; 18. upper laminal cells, x640; 19. leaf apex, xl70. B. hornschuchiana (20-27):
20. habit, xl; 21. habit, x 10; 22. leaves, x40; 23. upper leaf, x40; 24. leaf in cross section, x200; 25. basal
cells near costa (left side), x 170; 26. upper laminal cells, x640; 27. leaf apex, x 170. (1-10, Sim 9598; 11-19,
Magill 4266; 20-27, Schelpe 7802a).
240
Pottiaceae
Map 92. — • Barbula acutata
x Barbula afrofontana
of marginal or costal cells into differentiated photo-
synthetic tissues is an adaptation to harsh environ-
ments, expressed through convergent evolution by
several genera in the Pottiaceae, e.g. Tortula, Barbula ,
Crossidium, Pterygoneurum, Aloina and Acaulon.
Because of the apparently intergeneric configuration
of Pseudocrossidium, the species referrable there, have
been placed under various genera until further studies
can resolve the problems.
3. Barbula hornschuchiana Schultz in
Flora, Jena 5 (Syll.) : 35 (1822); Broth, in
Natiirl. PflFam. 10: 278 (1924); Sim, Bryo. S.
Afr. 239 (1926); Smith, Moss FI. Brit. Irel.
252 (1978). Type: Europe.
Plants small, caespitose, dark green to
reddish brown; terricolous. Stems 2-6 mm
tall, infrequently branched; in section with
small central strand, inner cortical cells
lax, outer cortical cells in 1-2 rows, stereids,
dark red. Leaves spirally twisted around
stem dry, erect-spreading wet, ovate-acumi-
nate to triangular, 0,8-1, 5 mm long; apex
acuminate, apiculate to cuspidate; margins
spirally revolute, frequently involving com-
plete upper lamina. Costa short-excurrent,
broad, ventral superficial cells quadrate to
rounded, sparsely papillose, dorsal super-
ficial cells short-rectangular, smooth; in
section flattened to reniform, guide cells
4-6, ventral stereid band in 1-2 rows, cells
infrequently substereids or only incrassate,
ventral surface cells larger, incrassate, papil-
lose, dorsal stereid band 4-5 cells thick, dorsal
Map 93. — • Barbula eubryum
X Barbula hornschuchiana
surface cells frequently substereids, smooth.
Upper laminal cells rounded-quadrate, incras-
sate, sharply mammillose or rarely with 4-6
low, fine, C-shaped papillae over lumen;
basal cells larger, quadrate to rectangular,
smooth.
Sporophyte not known in Africa. Middle
Eastern and European specimens have:
perichaetial leaves longer; seta short, orange;
capsule narrowly elliptical, weakly curved,
dark brown; operculum long-rostrate; spores
8-10 pm. Fig. 69: 20-27.
Barbula hornschuchiana is reported from north-
ern and Southern Africa, Europe and the Middle
East. In Southern Africa, the species has been col-
lected in the semi-arid shrublands of the southern
and western Cape. Map 93.
Vouchers: Magill 6080; Schelpe 7732.
The species is recognized by its ovate-acuminate
leaves with broad costa and strongly revolute margins.
The species was recently transferred to Pseudocros-
sidium by Zander (1979). The Southern African
specimens do not show specialization of marginal
or costal tissues characteristic of the genus.
4. Barbula calycina Schwaegr., Sp.
Muse. Suppl. 2: 63 (1823); Broth, in Natiirl.
PflFam. 10: 280 (1924). Type: Australia,
La Billardiere s.n.
Tortella calycina (Schwaegr.) Dix. in N. Zeal. Inst.
Bull. 3: 124 (1924); Scott & Stone, Moss. S. Aust.
200 (1976).
Tortula flexuosa Hook., Musci Exot. 2: 125 (1819),
horn, illeg., non Brid. (1806). Barbula flexuosa (Hook.)
POTTIACEAE
241
Schultz in Nova Acta Acad. Leop. Carol. 11: 208
(1823). Barbula hookeri Steud. in Nom. Bot. 2: 72
(1824). Type: Cape, Menzies s.n., 1791 (BM, holo.!).
Plants medium-sized, loosely caespitose,
yellow-green above, reddish brown below;
terricolous. Stems 10-20 mm tall, branching
by subperichaetial innovations, occasionally
with white tomentum below; in section sub-
round, central strand present, small, inner
cortical cells large, thin-walled, outer cortical
cells in 1-2 rows, stereids, reddish. Leaves
larger above, incurved dry, erect-spreading
to widespreading above erect base wet;
lanceolate to ovate-ligulate, 2-3 mm long;
apex acute to obtuse, mucronate; base
oblong to weakly obovate, appressed to
stem; margins plane, entire. Costa short-
excurrent, ventral superficial cells quadrate,
papillose, dorsal superficial cells quadrate to
short-rectangular, incrassate, papillose; in
section semicircular to reniform, guide cells
6, ventral stereid band strong, 2-4 cells
thick, ventral surface cells larger, thin-walled,
papillose, dorsal stereid band strong, 2-4
cells thick, dorsal surface cells incrassate,
papillose. Upper laminal cells subhexagonal
to quadrate, weakly incrassate, with 4-6 large,
simple, well-spaced papillae over lumen, not
obscuring cells; upper basal cells rectangular
to linear, yellowish, seriate papillose, ex-
tending down along costa, occasionally to
base, lower basal cells oblong-hexagonal to
rectangular, hyaline to yellowish, smooth,
thin-walled.
Dioicous. Perichaetial leaves highly dif-
ferentiated, lower leaves abruptly short-
subulate above elliptical, sheathing base,
cells of subula papillose, upper leaves spirally
sheathing seta, 3-5 mm long, irregularly
serrate at apex; leaf cells fusiform, sinuate.
Seta 20-50 mm long, yellowish to reddish;
capsule cylindrical, 2,5 mm long, with very
short neck, yellowish red; peristome 1,6-1, 8
mm long, cleft to near base, teeth linear,
reddish, strongly papillose, twisted 1^ turns
counter-clockwise, united below into short
basal membrane, 50 //m high, papillose;
operculum long-conic, 2,3 mm long, cells
twisted counter-clockwise; calyptra long-
cucullate; spores round, 8-10 pm, smooth.
Fig. 70: 1-9.
Barbula calycina is common in flat, sandy areas
of Australia, New Zealand, southeast Asia and
South America. The single South African collection
was reported from the fynbos biome of the south-
western Cape. Map 94.
Voucher: Menzies s.n., 1791 (BM!).
The Menzies collection is identical to plants
examined from Australia. The species is rather com-
mon in parts of Australia (Scott, pers. comm.) but
was collected only once from South Africa. Since
Menzies’s travels included both South Africa and
Australia in 1791, an error in labelling may have
occurred. The inclusion of B. calycina in the Flora is
provisional until additional material is collected.
Vegetatively B. calycina is closely related to B.
microcalycina, although the two species differ in leaf
shape, as well as several sporophytic characters.
5. Barbula microcalycina Magill, sp.
nov., habitu et cellulis foliorum basilibus
distinctis singularibusque B. calycina Schwaegr.
similis, sed forma foliorum, seta breviore,
peristomio brevi recto, operculo brevissimo et
sporibus majoribus differt.
Type: Natal, Cathedral Peak, Upper
Indumeni Forest, Magill 5706 (PRE, holo.;
C; H; MO; NY).
Plants small to medium, loosely caespi-
tose, yellow-green to light green, glaucous-
green dry, brownish below; terricolous or
saxicolous. Stems 8-20 mm high, thin,
branching by subperichaetial innovations;
in section round, central strand present,
small, frequently collapsed, inner cortical
cells large, thickened, outer cortical cells in
1-3 rows, stereids, reddish. Leaves about
equal throughout, incurved dry, widely
spreading to squarrose above clasping base
wet, weakly keeled; broadly lanceolate to
ligulate above obovate base, 1,5-2, 5 mm
long; apex broadly acute to obtuse, mucro-
nate; base sheathing; margins plane, entire.
Costa short-excurrent, strong, ventral super-
ficial cells quadrate to short-rectangular,
papillose, dorsal superficial cells quadrate to
short-rectangular, incrassate, papillose; in
section subround, guide cells 4, ventral
stereid band strong, in rounded group, 4 cells
thick, ventral surface cells similar to laminal
cells, papillose, dorsal stereid band strong,
3-4 cells thick, dorsal surface cells incras-
sate, papillose. Upper laminal cells rounded,
quadrate, incrassate, papillae 4-6 over cell,
low, simple or sometimes C-shaped, obscur-
ing cells; basal cells highly differentiated,
242
POTTIACEAE
upper basal cells rectangular or fusiform,
seriate papillose, lower basalcells rectangular,
hyaline, thin-walled, smooth.
Dioicous. Perichaetia terminal, leaves
differentiated, 1,2-1, 5 mm long; lanceolate
to acuminate above long-oblong, sheathing
base, papillose above. Seta 10-15 mm long,
yellow; capsule cylindrical, 1 ,8 mm long, with
very short neck, yellowish red; peristome
present, teeth irregularly cleft to base, 75 //m
long, straight, papillose, reddish yellow;
operculum short-conic, 0, 5-1,0 mm long,
cells weakly twisted counter-clockwise; calyp-
tra cucullate, to 3,5 mm long, twisted below,
spores round, 12-15 //m, smooth. Fig. 70:
10-19.
Fig. 70. — Barbula calycina (1-9): 1. habit, x 1 ; 2. habit, x 5; 3. leaf, x40; 4. leaf in cross section, x 170;
5. basal leaf cells, x 170; 6. upper laminal cells, x435; 7. perichaetial leaf, x20; 8. part od capsule mouth with
peristome teeth, x40; 9. operculum, X40. B. microcalycina (10-19): 10. habit, xl; 11. habit, x5; 12-13.
leaves, x40; 14. leaf in cross section, xl70; 15. basal leaf cells, xl70; 16. upper laminal cells, x640; 17.
perichaetial leaf, x 20; 18. operculum, X40; 19. part of capsule mouth with peristome teeth, X40. (1-9, Scott
775; 10-19, Smook 1418).
POTTIACEAE
243
Map 94. — • Barbula microcalycina
x Barbula calycina
Endemic to Southern Africa, B. microcalycina is
presently known only from the subalpine grasslands
of Lesotho and the Cathedral Peak and Giant’s
Castle areas of the Natal Drakensberg. Map 94.
Vouchers: Magill 4114, 5689, 5813; Pienaar 44;
Smook 1118, 1418; Van Rooy 4.
This species is similar to B. calycina, but differs
in the ligulate leaves with obovate, sheathing bases,
shorter seta, short, straight peristome and much
shorter operculum. It can be separated from other
species by its highly differentiated basal leaf cells.
6. Barbula indica {Hook.) Spreng. in
Steud., Nom. Bot. 2: 72 (1824); Saito in J.
Hattori bot. Lab. 39: 488 (1975); Sim,
Bryo. S. Afr. 236 (1926). Syntypes: India
Orientalis, Rottler s.n.; Horti Botanici Cal-
cuttae, Indiae Orientalis, Wallich s.n.
Tortula indica Hook., Musci Exot. 2: 135 (1819).
Semibarbula indica (Hook.) Herz. ex Hilp. in Beih.
bot. Zbl. 50: 626(1933).
Trichostomum orientalis Web. in Arch., Syst.
Naturgesch. 1: 129 (1804). Barbula orientalis (Web.)
Broth, in Natiirl. PflFam. 1: 403 (1902), horn, illeg.,
non B. orientalis Brid., Mant. Muse. 3: 93 (1819).
Semibarbula orientalis (Web.) Wijk & Marg. in Taxon
8: 75 (1959). Type: Asia, India Orientalis, Klein s.n.
Hymenostomum opacum Wag. & Dix. in Trans. R.
Soc. S. Afr. 4: 3 (1914). Type: Natal, Van Reenen,
Wager s.n. (PRE, holo. !).
Barbula natalensis C. Mull, in Hedwigia 38: 106
(1899). Type: Natal, Port Natal (Durban), Rehmann
s.n., vide Dixon in S. Afr. J. Sci. 18: 313 (1922).
Plants small, loosely caespitose, light
green; terricolous. Stems 2-8 mm tall,
simple; in section round, central strand
present, inner cortical cells lax, becoming
smaller toward margin, incrassate, outer
cortical cells in 1-2 rows, substereids, reddish
brown. Leaves crisped dry, spreading wet;
lanceolate to ovate-ligulate, 1,0-1, 8 mm
long; apex obtuse, apiculate; margins plane
to infrequently recurved below, entire. Costa
short-excurrent, ventral superficial cells rec-
tangular, smooth, dorsal superficial cells
rectangular, strongly prorate; in section
subround, guide cells 2-4, ventral stereid
band small, 1-2 cells thick, ventral surface
cells substereids or only incrassate; dorsal
stereid band large, 4-5 cells thick, dorsal
surface cells substereids, outer walls strongly
thickened, lumen frequently lunate. Upper
laminal cells quadrate or some hexagonal,
somewhat thickened, with 3-4 large, C-
shaped papillae over lumen; basal cells
oblong, elongate, hyaline, smooth. Gemmae
axillary or on axillary rhizoids, multicellular,
obovoid, 70-100 pm, brownish.
Dioicous. Perichaetia terminal; leaves
to 2 mm long, weakly differentiated. Seta
4-6 mm long, yellowish; capsules short-
cylindrical, 0,8-1 ,2 mm long, reddish yellow;
peristome erect or weakly twisted, fragile,
teeth linear, papillose, red-brown; operculum
conical, 0,4-0, 6 mm long; spores round,
10-14 pm, smooth. Fig. 71: 1—12.
Barbula indica occurs in Southern and central
Africa, the Indian subcontinent, southern and south-
east Asia and Japan. In Southern Africa the species
grows on soil in grasslands of Transvaal, Lesotho,
Natal and Transkei. Map 95.
Map 95.— • Barbula indica
x Barbula rehmannii
244
P0TT1ACEAE
POTTIACEAE
245
Vouchers: Cholno/cy 653; Magill 3615; Sim
10091 ; Stir ton 6948.
The strongly prorate dorsal costal cells and
numerous obovoid, multicellular gemmae charac-
terize B. indica. Specimens are only infrequently
collected with sporophytes.
7. Barbula rehmannii C. Mull, in Hedwi-
gia 38: 106 (1899); Broth, in Natiirl. PflFam.
10: 280 (1924). Type: Cape, Touws River,
Rehmann 101 (PRE!).
Plants small, caespitose, yellow-green;
terricolous. Stems 2-5 mm high, infrequently
branched; in section round, central strand
present, inner cortical cells lax, outer corti-
cal cells in 1-2 rows, smaller, incrassate,
red-brown. Leaves contorted dry, patent
wet; oval-ligulate to lanceolate, 1,0-1, 5
mm long; apex obtuse to obtusely acute;
margins plane to reflexed in base, entire.
Costa ending several cells below apex,
ventral superficial cells rectangular, sparsely
papillose, dorsal superficial cells long-rec-
tangular, incrassate, seriate papillose; in
section subreniform, guide cells 4, ventral
stereid band 1-2 cells thick, ventral surface
cells not differentiated, dorsal stereid band
3-4 cells thick, dorsal surface cells not
differentiated. Upper laminal cells quadrate to
subquadrate, incrassate, strongly papillose,
papillae C-shaped, 4-5 per cell; basal cells
larger, rectangular, papillose almost to base.
Dioicous. Perichaetia terminal; leaves
from very broad sheathing base, abruptly
narrow-acuminate, 1 ,0 mm long. Seta long,
to 20 mm, red-brown; capsule cylindrical,
1, 5-2,0 mm long, slightly curved, red-
brown ; peristome fragile, yellowish, only
short basal membrane seen; other parts not
seen. Fig. 71 : 13-19.
Endemic to Southern Africa, B. rehmannii has
been collected in fynbos of the southern and south-
western Cape and grasslands of the eastern Transvaal.
Map 95.
Voucher: Magill 3240.
Similar to B. indica in habit, B. rehmannii is
identified by its subpercurrent costa, papillose dorsal
costal cells and very long seta.
8. Barbula eubryum C. Miill. in Flora,
Jena 62: 379 (1879). Type: Tropical East
Africa, Ukamba Kitui, Hildebrandt s.n.,
1877.
Tortula eubryum (C. Miill.) Broth, in Engler,
Pflanzenw. Ost.-Afr. C: 69 (1916); Sim, Bryo. S.
Afr. 228 (1926).
Plants medium-sized, forming loose
tufts, yellow-green, blackish below; terri-
colous or saxicolous. Stems 10-15 mm tall,
infrequently branched, tomentose below;
in section round, central strand small or
rarely absent in some plants, inner cortical
cells large, outer cortical cells in 1-2 rows,
smaller, incrassate, red-yellow. Leaves spirally
twisted around stem dry, patent wet, rugose
to rugulose, keeled; ligulate to broadly
lanceolate, 3, 5-4,0 mm long; apex obtuse to
broadly acute, mucronate to apiculate ;
margins plane to reflexed below, entire.
Costa short-excurrent, ventral superficial
cells rectangular, thin-walled, smooth, dorsal
superficial cells elongate, smooth, incrassate;
in section reniform to subround, guide cells
3-4, ventral stereid band small, in one row
over central guide cells, ventral surface cells
incrassate, dorsal stereid band large, 4-5
cells thick, dorsal surface cells substereids.
Upper laminal cells hexagonal to quadrate,
papillose with 4-6 low papillae over lumen;
basal leaf cells large, rectangular, smooth,
becoming narrowly rectangular toward
margin.
Sporophytes not seen on Southern
African plants. Setae in Ugandan plants are
10-12 mm long and capsules cylindrical,
Fig. 71. — Barbula indica (1-12): 1. habit, x 1 ; 2. habit, x 1 ; 3. habit, x 5; 4-5. leaves, x 30; 6. leaf in cross
section, x435; 7. basal leaf cells (left side), x250; 8. upper laminal cells, x640; 9. leaf apex, dorsal surface
(papillae partly shown), x250; 10. rhizoid gemmae, x40; 11. axillary gemmae, x300; 12. capsule mouth with
peristome, x30. B. rehmannii (13-19): 13. habit, xl; 14. habit, x 5; 15. leaves, x30; 16. leaf in cross section,
x435; 17. basal leaf cells (right side), x250; 18. upper laminal cells, x640; 19. leaf apex, x250. B. eubryum
(20-27): 20. habit, wet, x 1 ; 21. habit, dry, x 1 ; 22. habit, x 5; 23. leaves, X 15; 24. leaf in cross section, x 190;
25. cells at leaf base (right side), x90; 26. upper laminal cells, x300; 27. leaf apex (papillae not shown), x90.
B. afrofontana (28-34): 28. habit, x 1 ; 29. habit, x 2; 30. leaves, x 20; 31. leaf in cross section, x 320; 32. basal
leaf cells (left side), x 170; 33. upper laminal cells, x 435; 34. leaf apex, X 170. (1-3, Stirton 6958; 4-12, Cholnoky
447; 13-19, Rehmann 101 ; 20-27, Magill 3580; 28-34, Rehmann 82).
246
POTTIACEAE
2, 5-3,0 mm long. The Ugandan plants are
frequently twice as tall as Southern African
specimens. Fig. 71 : 20-27.
Barbula eubryum is known from central and
Southern Africa. In the south the species is found on
soil or rock in grassland and woodland communities
of central and eastern Transvaal, Swaziland, Zululand
and Natal. Map 93.
Vouchers: Harrison 14; Magill 3571, 3580; Van
Vuuren 1742.
The large plants with broad, rugose leaves are
very distinctive and characterize the species. The size
and habit of the plants suggest a relationship to
Tortula, but the presence of a ventral stereid band in
the costa indicates that the species is correctly placed
in Barbula.
9. Barbula afrofontana (C. Mull.) Broth.
in Natiirl. PflFam. 10: 280 (1924); Sim, Bryo.
S. Afr. 235 (1926). Type: Natal, Van Reenens
Pass, Rehmann 82 (PRE!).
Trichostomum afrofontanum C. Miill. in Hedwigia
38: 99 (1899). Hydrogonium afrofontanum (C. Mull.)
Hilp. in Beih. bot. Zbl. 50: 632 (1933).
Plants long, light to dark green; aquatic
to semi-aquatic, tufaceous saxicolous (didy-
modontoliths). Stems flaccid, to 50 mm long,
green to blackish; in section round, central
strand very small, of 4-5 cells, inner cortical
cells lax, outer cortical cells in 2-3 rows,
smaller, incrassate. Leaves distant, lax,
spreading, frequently lime-encrusted; broadly
lanceolate to triangular-lanceolate, 2-3 mm
long; apex obtuse; margins plane, entire,
frequently indistinctly bordered. Costa ending
2-5 cells below apex or percurrent, ventral
and dorsal superficial cells rectangular,
smooth; in section subround, guide cells 4,
ventral stereid band small, 1-2 cells thick
over central guide cells, ventral surface cells
larger, incrassate, dorsal stereid band 3-4
cells thick, dorsal surface cells larger, incras-
sate. Upper laminal cells hexagonal, quadrate
or short-rectangular, superficially bulging,
smooth or occasionally with 1-2 indistinct
papillae in upper leaf; marginal cells in 4-5
rows, narrower, weakly incrassate, rectangu-
lar to quadrate; in section frequently forming
indistinct, elliptical border; basal cells elon-
gate-rectangular, becoming somewhat nar-
rower toward margin. Gemmae axillary,
multicellular, fusiform, brown or green.
Sporophyte unknown. Fig. 71 : 28-34.
Barbula afrofontana is presently known only
from eastern and Southern Africa. In the Flora area,
the species occurs in or around waterfalls or seepage
areas in South West Africa/Namibia, eastern Trans-
vaal, Natal, Transkei and eastern Cape. Map 92.
Vouchers: Cholnoky 853; Giess 8993; Phelan &
Smook 70; Van Rooy 283.
The species is identified by its semi-aquatic
nature, flaccid habit, subpercurrent costa, smooth
leaf cells and weak leaf border. The border on leaf
margins is rarely obvious, since it consists of slightly
enlarged and thickened marginal cells.
Insufficiently Known Species
Barbula anoectangiacea C. Miill. in Hedwigia 38:
105 (1899). Type: Cape, Somerset East, Boschberg,
MacOwan s.n. The type has not been seen. Brotherus
in Natiirl. PflFam. 1: 403 (1902) suggested that this
was a species of Ceratodon.
Barbula flexicaulis C. Miill. in Hedwigia 38: 107
(1899). Type: Orange Free State, Taaiboschkranz,
Rehmann s.n., 1875. The type has not been seen.
Brotherus in Natiirl. PflFam. 1 : 411 (1902) suggested,
after examining the specimen, that it was a Schistidium.
Sim (1926) referred the species to Grimmia apocarpa,
although he had not seen the type.
Barbula perlinearis C. Miill. in Hedwigia 38: 107
(1899). Type: Transvaal, Lydenburg, Wilms s.n.,
Feb. 1888. The type has not been seen. Sim’s (1926)
treatment of this species under Didymodon xantho-
carpus (C. Mull.) Magill (as Barbula xanthocarpa C.
Mull.), does not appear correct, since the description
and known distribution of that species do not corres-
pond with those of B. perlinearis.
Barbula pertorquata C. Mull, in Hedwigia 38: 109
(1899). Type: Cape, Cape Town, Rehmann s.n., 1875.
The type has not been seen. Sim (1926) refers the
species to Didymodon xanthocarpus (C. Miill.) Magill
(as Barbula xanthocarpa C. Miill.).
5. BRYOERYTHROPHYLLUM
Bryoerythrophyllum Chen in Hedwigia 80: 4 (1941); Zander in Bryologist 81: 541 (1978).
Type species: B. recurvirostrum (Hedw.) Chen.
Plants small to medium, loosely caespitose or in dense tufts, reddish; terricolous or
saxicolous. Stems erect, branching by subperichaetial innovations; central strand present.
Leaves distant or rather dense; ligulate to narrowly lanceolate; apex acute, mucronate; base
POTTIACEAE
247
Fig. 72. — Bryoerythrophyllum jamesonii (1-10):
1. habit, x 1 ; 2. habit, x3; 3. habit, x 10; 4. stem in
cross section, xl30; 5. leaves, x40; 6. leaf in cross
section, x640; 7. cells at leaf baes (left side), x 170;
8. upper laminal cells, x640; 9. leaf apex, x 170; 10.
part of capsule mouth with peristome teeth and
spores, x 170. B. recurvirostrum (11-18): 11. habit,
xl; 12. habit, xl; 13. habit xlO; 14. stem in cross
section, x 100; 15. leaf, x 30; 16. leaf in cross section,
x230; 17. upper laminal cells at margin (dorsal view),
X320; 18. part of capsule mouth with peristome teeth
and spores, x 140. (1-10, Magill 4329; 11-18, Magill
4446).
248
POTTIACEAE
clasping or appressed; margins plane to revolute, entire or irregularly toothed near apex.
Costa short-excurrent; with dorsal and ventral stereid bands. Laminal cells quadrate to short-
rectangular, papillose; basal cells larger, rectangular, smooth.
Paroicous to dioicous. Capsule long-exserted, short-cylindrical; peristome fragile, teeth
linear, spiculate; operculum short-conic to rostrate; calyptra cucullate, smooth; spores
subround, smooth.
Bryoerythrophyllum, with 17 species, is best represented in Eurasia. Only 7 of these species are recognized
in the Americas and only the two most widely distributed species are known from Africa and Australia. In
Southern Africa the genus is found at upper elevations of the Drakensberg and forests of Natal and eastern
Transvaal.
1 Leaf margins plane above mid-leaf 1. B.jamesonii
1 Leaf margins revolute to near apex 2. B. recurvirostrum
1. Bryoerythrophyllum jamesonii ( Tayl .)
Crum in Svensk bot. Tidskr. 51: 200 (1957);
Zander in Bryologist 81: 549 (1978). Type:
Ecuador, Pichincha, Quito, Jameson 88
(NY).
Barbula jamesonii Tayl. in J. Bot., Lond. 5: 48
(1846).
Didymodon afrorubellus Broth. & Wag. ex Dix. in
Trans. R. Soc. S. Afr. 8: 194 ( 1 920) ; Broth, in Natiirl.
PflFam. 10: 273 (1924); Sim, Bryo. S. Afr. 248 (1926).
Bryoerythrophyllum afrorubellum (Broth. & Wag.) De
Sloover in Bull. Jard. nat. bot. Belg. 49: 398 (1979).
Type: Natal, Van Reenens Pass, Wager 79 (PRE,
holo.!; BM!; H).
Plants small to medium, in dense or
loose tufts, dark green, reddish brown
below; terricolous or saxicolous. Stems
erect to inclined, 2-15 mm high, branching;
in section angular, central strand present,
inner cortical cells large, outer cortical cells
in 1-3 rows, stereids or substereids. Leaves
incurved, contorted dry, erect-spreading wet,
somewhat distant; ligulate to oblong-lanceo-
late, 1,0-1, 5 mm long; apex acute, mucro-
nate; base appressed to stem; margins fre-
quently recurved just above base, plane
above, entire or irregularly dentate by smooth,
yellowish marginal cells. Costa short-excur-
rent, ventral superficial cells quadrate to
short-rectangular, papillose, dorsal super-
ficial cells elongate, incrassate, sparsely
papillose; in section reniform, guide cells 4,
ventral stereid or substereid band 1 cell
thick, ventral surface cells incrassate, papil-
lose, dorsal stereid band 2-A cells thick,
dorsal surface cells undifferentiated. Upper
laminal cells quadrate to rounded-quadrate,
with 4-6 low, blunt papillae over lumen;
basal cells larger, well defined, rectangular,
smooth, reddish yellow.
Dioicous. Male plants somewhat smaller,
perigonia terminal; leaves with broadly
sheathing bases. Perichaetia terminal; leaves
oblong-lanceolate, to 2 mm long. Seta
0,7-1, 2 mm long, reddish; capsule cylindri-
cal, 1,0- 1,2 mm long, reddish brown;
peristome fragile, teeth erect, linear, fre-
quently perforated below, spiculate, reddish
yellow; operculum conical, 0,6 mm long;
spores round, 17-20//m, granulate to smooth.
Fig. 72: 1-10.
New to Southern Africa, B. jamesonii is known
from Scotland, British Columbia, Central and South
America, India, China, Australia and New Zealand.
In Southern Africa, B. jamesonii is most frequently
collected in the mountain grasslands of Lesotho,
eastern Orange Free State and western Natal. The
species is sporadically collected in forests or sheltered
kloofs of the Transvaal and central Natal and several
specimens are also known from the mountains of the
southwestern Cape. Map 96.
Vouchers: Brenan M2847; Cholnoky 17; Magill
4329, 4510,5133.
Bryoerythrophyllum jamesonii is generally defined
by the reddish tint of the plants, small leaves with
plane margins, highly differentiated basal cells and the
few apical teeth. A few specimens have been seen that
lack the reddish colouration and/or apical teeth on
the leaves, but have been identified by a combination
of other characters including habit, leaf shape, basal
leaf cells and sporophyte.
2. Bryoerythrophyllum recurvirostrum
(Hedw.) Chen in Hedwigia 80: 5 (1941);
Zander in Bryologist 81: 542 (1978). Type:
Europe.
Weissia recurvirostris Hedw., Spec. Muse. 71
(1801).
Plants small to medium, forming dense
tufts or cushions, brownish green above,
reddish below; terricolous or saxicolous.
Stems erect, 0,5-20,0 mm high, branching;
POTTIACEAE
249
in section with large central strand, inner
cortical cells lax, outer cortical cells in 1-2
rows, small, incrassate, red. Leaves con-
torted dry, erect-spreading wet, dense; lanceo-
late to linear-lanceolate above ovate to oblong
base, 2, 0-3, 5 mm long; apex acute, mucro-
nate; margins revolute to apex, entire or fre-
quently with a few yellowish teeth at apex.
Costa short-excurrent, ventral superficial cells
quadrate, incrassate, papillose, dorsal superfi-
cial cells elongate, incrassate, papillose; in
section suboval, guide cells 2-4(-6), lateral
guide cells sometimes smaller, ventral stereid
band small, 2 cells thick, surface cells larger, in-
crassate, papillose, dorsal stereid band 3-4 cells
thick, surface cells slightly larger, outer walls
strongly incrassate, lumens lunate. Upper
laminal cells quadrate to rectangular, rarely a
few cells transversely rectangular, papillose,
3-4 low, C-shaped papillae covering cell;
marginal cells papillose ; basal cells rectangu-
lar, smooth, red-brown.
Paroicous. Perichaetial leaves with wide,
sheathing base. Seta straight, 5-7 mm long,
red-brown; capsule short-cylindrical, 1,0-
1,5 mm long, dark red when mature, light
brown when old; peristome teeth erect,
linear, cleft or perforated above middle,
spiculate, yellowish; operculum obliquely
rostrate, to 0,6 mm long; spores subround,
17-20 pm, smooth. Fig. 72: 11-18.
Map 96. — • Bryoerythrophyllum jamesonii
X Bryoerythrophyllum recurvirostrum
Widespread i.i the Northern Hemisphere, B.
recurvirostrum is uncommon or rare in Africa (Bizot
et at., 1978) Australia and New Zealand and unknown
in South America. In Southern Africa the species is
found in the subalpine grasslands of Lesotho and
Natal, and forest in the eastern Cape. Map 96.
Vouchers: Jacot Guillarmod 4583; Magill 4195,
4446.
The species is identified by its reddish colouration,
lanceolate leaves with recurved margins, apical teeth
and strongly differentiated basal leaf cells. The size of
the plants, leaf shape and recurved margins distin-
guish B. recurvirostrum from B. jamesonii.
Subfamily Trichostgmoideae
Plants small to large, in loose tufts or scattered individuals; terricolous, saxicolous or
corticolous. Stems erect; central strand present or absent. Leaves generally narrow, linear to
lanceolate or oblong to elliptical, rarely narrowly spatulate; margins plane to involute. Costa
subpercurrent to short-excurrent, rarely long-excurrent; in section with dorsal and ventral
stereid bands. Upper laminal cells small, quadrate, papillose; junction with basal cells straight
across leaf or basal cells extending up margins forming distinct V-shaped junction with laminal
cells. Gemmae absent.
Capsules cylindrical; peristome present or absent, teeth short, erect or long-filamentous,
generally twisted; operculum rostrate; calyptra cucullate, long.
Key to Genera of Subfamily Trichostomoideae
1 Leaves bistratose above base:
2 Leaves bistratose juxtacostally; costa subpercurrent 3. Timmiella
2 Upper lamina bistratose throughout; costa long-excurrent 4. Tortella
250
POTTIACEAE
1 Leaves unistratose above base :
3 Leaves broadly lanceolate to Ungulate; capsules exserted, cleistocarpic or immersed,
stegocarpic:
4 Capsules cleistocarpic, longitudinally ridged dry; leaf margins plane; costa broad,
percurrent 1 . Tetrapterum
4 Capsules stegocarpic, immersed, gymnostomous; leaf margins erect, incurved
above; costa mucronate 2. Phasconica
3 Leaves linear-lanceolate to oblong; capsules long-exserted, stegocarpic:
5 Basal hyaline cells extending up margins in lower -J— } of leaf, forming distinct
V-shaped area in leaf base; peristome long, twisted 4. Tortella
5 Basal hyaline cells not forming V-shaped area or only weakly so, junction with
upper laminal cells ± straight across leaf ; peristome present or absent, generally
short, not twisted:
6 Central strand of stem absent 5. Oxystegus
6 Central strand of stem present :
7 Leaves narrowly spathulate to elliptical; margins plane 6. Trichostomum
7 Leaves linear-lanceolate to oblong; margins incurved to involute 7. Weissia
1. TETRAPTERUM
Tetrapterum Hampe ex Jaeg. in Verh. St Gall, naturw. Ges. 1868-69: 85 (1869); Broth, in
Naturl. PflFam. 10: 252 (1924); Sim, Bryo. S. Afr. 254 (1926). Type species: T. australe
Hampe.
Plants small, light or yellow-green; terricolous. Stems erect; central strand present.
Leaves broadly lanceolate above oval or oblong base; margins plane, entire. Costa strong,
short-excurrent, broad below. Upper laminal cells quadrate, with 4-6 O-shaped papillae over
lumen.
Seta short; capsule exserted, cleistocarpic, urn longitudinally ridged dry, bluntly beaked.
Eleven species are currently recognized for Tetrapterum. The genus is known as far north as Central
America, but is mainly restricted to southern South America, Southern Africa and Australia. The genus is
recognized by broad leaves with plane margins, the presence of dorsal and ventral stereid bands in the costa,
and cleistocarpic capsules.
Tetrapterum tetragonum {Hook.) An-
drews in Bryologist 48: 191 (1945); Sim,
Bryo. S. Afr. 254 (1926). Type: Cape, Tiger
Berg, Mund s.n.
Phascum tetragonum Hook, in Bot. Misc. 1: 124
(1829). Astomum tetragonum (Hook.) Broth, in
Naturl. PflFam. 1 : 385 (1902).
Plants small, loosely caespitose, yellow-
green; terricolous. Stem 2-4 mm tall, rarely
branched; in section round, central strand
large, cortical cells in 5-6 rows, thin-walled,
smaller toward margin. Leaves spirally
twisted around stem dry, erect-spreading
wet; lanceolate, 1,5-1, 8 mm long; apex
acute, mucronate; base oblong or oval,
erect-sheathing; margins plane, entire. Costa
short-excurrent as 2-3 clear, smooth cells,
ventral and dorsal superficial cells quadrate,
papillose; in section reniform, guide cells 4-8,
ventral stereid band 2-3 cells thick, ventral
surface cells larger, quadrate, incrassate,
papillose, dorsal stereid band 3-4 cells thick,
dorsal surface cells quadrate, incrassate,
papillose. Upper laminal cells quadrate,
thin-walled, papillae low, 4-6 over lumen,
O-shaped; basal cells rectangular, thin-
walled, hyaline, smooth.
Autoicous. Perigonia lateral on short
branches; perichaetia terminal, leaves undif-
ferentiated. Seta 1 mm long, yellow; capsule
POTTIACEAE
251
Map 97. — • Tetrapterum tetragonum
X Phasconica tisserantii
cleistocarpic, urn elliptical, bluntly beaked,
with strong longitudinal ridges dry, ±
quadrate in median transverse section, 1 , 5
mm long, red-yellow; exothecial cells rec-
tangular, thin-walled; stomata phaneropore,
present at base of urn; calyptra cucullate,
obliquely beaked; spores round, 20-25
pm, papillose. Fig. 73: 1-9.
Endemic to Southern Africa, Tetrapterum tetra-
gonum is infrequently collected in dry shrublands of
the southwestern and northwestern Cape. Map 97.
Vouchers: Goldblatt 619; Magill & Schelpe 3861,
3870; Schelpe 7628.
The leaf shape and costal anatomy are quite
distinctive and help to identify sterile specimens. The
cleistocarpic, orange capsules, produced in abun-
dance, are strongly ridged when dry and are more or
less rectangular wet. The distinctive capsules charac-
terize specimens of Tetrapterum.
2. PHASCONICA
Phasconica C. Mull, in Linnaea 43: 438 (1882). Type species: P. lorentzii C. Mull.
Plants small, gregarious, yellow-green; terricolous. Stems to 1 mm high; central strand
present. Leaves broadly lanceolate to Ungulate; margins erect to incurved above, entire.
Costa short-excurrent; with dorsal and ventral stereid bands. Upper laminal cells quadrate,
incrassate, papillose.
Seta very short ; capsule cyathiform, gymnostomous ; operculum plano-convex, abruptly
rostrate ; calyptra cucullate.
The genus Phasconica comprises three species from widely separated regions. The African species, P.
tisserantii P. Varde, is known from central and Southern Africa; P. lorentzii C. Mull, is known from South
America and P. balansae C. Mull, occurs in Oceania. Saito (1975) recently included the genus in the synonomy
of Weissia Hedw. without explanation. Judging from Muller’s descriptions only, Saito’s decision appears to be
correct ; however, I have not seen the latter two species. The African species is not a Weissia. It bears a strong
gametophytic resemblence to Trichostomum Bruch, but sporophytically, the two genera are very different.
The relationship between Phasconica, in the African sense, and Trichostomum is reminiscent of the relationship
between Weissia and the segregate genera Astomum Hampe and Hymenostomum R. Br.
Phasconica tisserantii P. Varde in Revue
bryol. lichen. 7: 231 (1935). Type: Central
African Republic, Bozoum, Tisserant s.n.
(PC).
Plants small, gregarious, yellow-green;
terricolous. Stems to 1 mm tall, simple; in
section round, central strand present, cortical
cells incrassate. Leaves larger above, in-
curved-contorted dry, spreading wet ; lanceo-
late to ligulate, 2-3 mm long; apex acute,
mucronate; base scarcely differentiated; mar-
gins plane to erect below, incurved above,
entire. Costa short-excurrent, ventral super-
ficial cells quadrate, papillose, dorsal super-
ficial cells rectangular, smooth; in section
semicircular, guide cells 4, ventral stereid
band 2 cells thick, ventral surface cells
similar to laminal cells, papillose, dorsal
stereid band 4-5 cells thick, dorsal surface
cells substereids. Upper laminal cells quadrate
to subquadrate, incrassate, with 4-6 low,
blunt papillae over lumen; basal cells rec-
tangular to long-hexagonal, lax, smooth.
Dioicous. Perigonia not seen; peri-
chaetia terminal, leaves ligulate, 3 mm long.
Seta erect, 0,3 mm long, yellowish; capsule
252
POTTIACEAE
POTTIACEAE
253
immersed, cyathiform, to 1 mm long, yellow-
ish, mouth red; stomata not seen; vaginula
oval-cylindrical, to 0,8 mm long; peristome
absent; operculum plano-convex, abruptly
rostrate, 0,6 mm in diameter, 0,5 mm high,
cells not twisted; spores round, (24-) 27-30
pm, strongly papillose. Fig. 73: 10—19.
The species is known from central Africa and
mountains of the southwestern Cape. Map 97.
Voucher: Gar side 6697.
The strongly costate leaves and immersed, oper-
culate capsule are quite distinctive in Southern
Africa and will identify P. tisserantii. Some doubt
exists as to the generic position of this species since
Saito (1975) considered Phasconica a synonym of
Weissia. This species is only very distantly related to
Weissia and may require a new combination if
Mueller’s types are indeed Weissia sens. lat. Game-
tophytically P. tisserantii resembles Trichostomum
brachydontium , but the large, immersed, cup-shaped,
gymnostomous capsule clearly distinguishes it from
that species.
3. TIMMIELLA
Timmiella ( De Not.) Limpr., Laubm. Deutschl. 1: 590 (1888); Broth, in Natlirl. PflFam. 10:
261 (1924). Type species: T. anomala (B.S.G.) Limpr.
Plants small, dark green; terricolous. Stems erect; central strand present. Leaves bistra-
tose; ligulate above oval base; in section bistratose juxtacostally, dorsal cells smaller; margins
irregularly serrate. Costa subpercurrent, dorsal and ventral stereid bands present. Laminal
cells quadrate, smooth or mammillose; basal cells lax.
Sporophyte terminal. Seta long, flexuose or straight; capsule cylindrical; peristome teeth
filamentous, straight or twisted.
The 15 species of Timmiella are scattered throughout the world with the exception of Australia and New
Zealand. Two species are widespread in the Northern Hemisphere; the other species are rather local in distribu-
tion. The genus is easily identified by its unusual leaf anatomy. In addition to T. pelindaba, 4 other species
occur in central or northern Africa.
Timmiella pelindaba Magill in Mem. bot.
Surv. S. Afr. 43: 3 (1979). Type: Transvaal,
Pelindaba, Bosman PRE-CHI 607 (PRE,
holo. ! ; H ; L ; MO ; NY).
Plants small, gregarious, dark green ;
terricolous. Stems to 5 mm high, unbranched ;
in section round, central strand small, inner
cortical cells lax, outer 2-3 rows incrassate,
reddish; axillary hairs of 6-8 cylindrical
cells, hyaline throughout. Leaves crisped
with involute margins dry, widely spreading
to recurved with plane margins wet; bistra-
tose juxtacostally; ligulate, 3, 5-4, 5 mm
long; apex acute; base oval; lamina frequent-
ly constricted above base; margins plane,
irregularly serrate in distal $, entire below.
Costa strong, subpercurrent, very wide below,
ventral superficial cells quadrate, smooth,
dorsal superficial cells quadrate, smooth; in
section round, guide cells 4, ventral stereid
band small, 1-2 cells thick, ventral surface
cells similar to ventral laminal cells, dorsal
stereid band 3-4 cells thick, dorsal surface
cells substereids, ± irregular. Upper laminal
cells quadrate, mammillose ventrally, smooth
dorsally; in section dorsal cells extending
about halfway to margin, c. 15 cells each
side of costa, half as high as ventral cells;
basal cells lax, oblong-hexagonal, thin-
walled.
Dioicous. Perichaetial leaves not dif-
ferentiated. Seta straight wet or dry, 12 mm
Fig. 73. — Tetrapterum tetragonum (1-9): 1. habit, x 1 ; 2. habit, dry, x 10; 3. habit, wet, x 10; 4. stem in
cross section, x 170; 5. leaves, x40; 6. leaf in cross section, x235; 7. cells at leaf base (left side), x 170; 8.
leaf apex (papillae partly shown), x 170; 9. spores, x270. Phasconica tisserantii (10-19): 10. habit, xl; 11.
habit, x 10; 12. stem in cross section, X 170; 13. leaves, x 30; 14. leaf in cross section, X 250; 15. cells at leaf
base (right side), xl70; 16. upper laminal cells (papillae partly shown), x640; 17. leaf apex (dorsal left side),
x 170; 18. calyptra, x 30; 19. capsule, x 30. (1-9, Giffen PRE-CH7723; 10-19, Garside 6697).
254
POTTIACEAE
long, reddish; capsule cylindrical, 2 mm long,
reddish yellow; exothecial cells oblong;
annulus well developed, persistent; peristome
teeth straight, long-filiform, 350-400 /zm
high, ± smooth, reddish yellow, connected at
base and forming a very short tube; oper-
culum long-rostrate, 1 mm long; calyptra
cucullate, 3,5 mm long; spores round,
12-15 /tm, essentially smooth. Fig. 74: 1-12.
Endemic to Southern Africa, this species is known
from kloofs in the grasslands of central Transvaal,
Lesotho and southeastern Orange Free State. Map 98.
Map 98. — • Oxystegus cylindricus
x Timmiella pelindaba
Vouchers: Smook 2319a; Van Rooy 597.
The species is near the central African T. cameru-
niae Broth., but differs in leaf length and shape,
plane margins and straightness and length of the seta.
It differs from other African species of Timmiella by
its dioicous condition, well developed annulus and
straight peristome teeth.
Fig. 74. — Timmiella pelindaba: 1. habit, wet,
xl; 2. habit, dry, xl; 3. habit, x 10; 4. stem in
cross section, xl25; 5-6. leaves, x20; 7. leaf in
cross section, x 260; 8. cells at leaf base (right side),
xl20; 9. leaf apex, xl70; 10. capsule, x 10; 11.
capsule mouth and part of peristome teeth, x90;
12. operculum, x 10. (1-12, Bosman PRE-CH1607).
POTTIACEAE
255
4. TORTELLA
Tortella ( Lindb .) Limpr., Laubm. Deutschl. 1: 520 (1888), nom. cons.; Saito in J. Hattori
bot. Lab. 39:440 (1975). Type species: T. caespitosa (Schwaegr.) Limpr.
Plants small to large, forming tufts or cushions; terricolous, saxicolous or corticolous.
Stem erect, central strand present or absent; axillary hairs of 10-20 cells, hyaline throughout.
Leaves incurved, variously contorted dry; linear-lanceolate to oblong; margins plane, entire.
Costa short to long-excurrent. Laminal cells quadrate to short-rectangular, incrassate, papil-
lose or surface sometimes strongly thickened to mammillose; basal cells strongly differentiated,
hyaline, smooth, extending up margins further than costa, forming distinct V-shaped interface
between basal and laminal cells.
Autoicous. Perigonia on short lateral branches; perichaetia terminal, leaves undifferen-
tiated. Capsule cylindrical; peristome long, twisted, teeth filiform above very short basal
membrane; operculum conic-rostrate; calyptra cucullate, long; spores papillose.
The genus Tortella contains 63 species evenly distributed throughout the world, with the exception of
Antarctica. A few of the species, i.e. T. humilis and T. fragilis, are almost cosmopolitan. The genus, although
not common, is found throughout Southern Africa and is recognized by its narrow leaves with plane margins,
casta with dorsal and ventral stereid bands, basal leaf cells extending well up leaf margins and long, twisted
peristome teeth above a short basal membrane.
1 Leaves long-subulate, rigid wet or dry, bistratose above; costa long-excurrent, fragile 3. T. fragilis
1 Leaves unistratose throughout; costa short-excurrent or mucronate:
2 Leaves oblong; apex broad, obtuse or rarely acute 2. T. humilis
2 Leaves lanceolate to linear; apex narrow, acute or occasionally obtuse 1.7. xanthocarpa
1. Tortella xanthocarpa (C. Mull.) Broth.
in Natiirl. PflFam. 1: 397 (1902). Type:
Cape, Saldanha Bay, Breutel s.n. (H-BR!).
Trichostomum xanthocarpum Schimp. ex C. Mull,
in Bot. Ztg 17: 229(1859).
Trichostomum tortuloides Sull. & Lesq. in Proc. Am.
Acad. Arts Sci. 4: 277 (1859); Sim, Bryo. S. Afr. 244
(1926). Tortella tortuloides (Sull. & Lesq.) Broth, in
Natiirl. PflFam. 1: 397 (1902). Type: Cape, Simons-
town, C. Wright s.n., 1853 (FH, holo.!).
Trichostomum rufisetum C. Mull, in Hedwigia 38:
99 (1899); Sim, Bryo. S. Afr. 243 (1926). Tortella
rufiseta (C. Mull.) Broth, in Natiirl. PflFam. 1 : 397
(1902). Type: Cape, Blanco, Rehmann sub 45 ( Reh -
mann 115, PRE!; BM!; vide Dix. & Gepp, 1923).
Trichostomum rehmannii Sim, Bryo. S. Afr. 244
(1926). Type: Transvaal, Pilgrims Rest, Maclea s.n.
( Rehmann 471; PRE, holo.!).
Plants small, caespitose, green to yellow-
green, brownish below; terricolous, corti-
colous or rarely saxicolous. Stems 1-5 mm
tall, occasionally branching; in section round,
central strand small, cortical cells incrassate,
smaller and reddish toward margin. Leaves
crowded above, contorted to incurved dry,
erect-spreading wet, somewhat fragile; linear
to lanceolate, 2, 5-4,0 (-5,0) mm long; apex
acute or occasionally some leaves obtuse,
frequently mucronate; base oval to oblong;
margin plane, entire or occasionally irregular
above through displaced cells. Costa short-
excurrent, ventral superficial cells quadrate,
papillose or mammillose, dorsal superficial
cells rectangular to linear, smooth; in section
subround, guide cells 4, large, ventral stereid
band 2-3 cells thick, ventral surface cells
similar to laminal cells, papillose or mammil-
lose, dorsal stereid band 2-5 cells thick,
dorsal surface cells not differentiated. Upper
laminal cells quadrate to short-rectangular,
weakly thickened, superficially bulging, papil-
lose to weakly papillose, 4-6 over lumen, low,
blunt or strongly thickened and becoming
mammillose, appearing smooth in surface
view; basal cells rectangular, lax, hyaline,
smooth, extending up margins, restricted to
lower i or extending to mid-leaf.
Autoicous. Perigonia on short lateral
branches; perichaetia terminal, leaves undif-
ferentiated. Seta 6-10 (-15) mm long, reddish
yellow; capsule cylindrical, 2 mm long;
peristome present, yellowish red, inserted at
mouth, teeth filiform above very short basal
membrane, 0,5 mm high, twisted counter-
clockwise; operculum rostrate, 0,8 mm long;
calyptra cucullate, covering urn ; spores
256
POTTIACEAE
round, 15-17 pm, weakly papillose. Fig. 75:
11-28.
Endemic to Southern Africa, T. xanthocarpa is
most frequently collected in forests or wooded areas
of the southern and western Cape and in moun-
tainous regions of the Orange Free State and Natal.
A few collections are also known from scattered
wooded areas in Swaziland, Transvaal, Zululand and
northern Cape. Map 99.
Map 99. — • Tortella xanthocarpa
X Tortella fragilis
Vouchers: Hilliard & Burtt 10115a; Lewis
12205; Magill 4083, 5523, 5605, 6104, 6291; Oliver
7081 ; Schelpe 4008; Van Rooy 374, 586.
The lanceolate leaves, acute apex, oval base and
short-excurrent costa will separate the typical ‘facies’
of T. xanthocarpa from the other species in Southern
Africa. I agree with Dixon (cf. Sim, 1926) that
Trichostomum rehmannii is a diminutive form of this
species and I also find T. tortuloides conspecific.
Tortella rufiseta is also included here as it represents a
robust form that is otherwise indistinguishable.
Many of the specimens, collected on tree bark in
forests of the Drakensberg and eastern Transvaal,
have longer, linear leaves with oblong bases and
narrow apices. These specimens may represent an
undescribed variety of T. xanthocarpa, although there
appears to be a fairly continuous gradation between
these linear-leaved specimens and the more common
lanceolate-leaved plants.
2. Tortella humilis ( Hedw .) Jenn., Man.
Moss. W. Pennsylv. 96 (1913); Redfearn in
Ann. Mo. bot. Gdn 59: 29 (1972); Crum,
Moss. Gt Lakes For. 104 (1972). Type:
North America, Pennsylvania.
Barbula humilis Hedw., Spec. Muse. 1 16 (1801).
Barbula caespitosa Schwaegr., Spec. Muse. Suppl.
1 : 120 (1811). Tortella caespitosa (Schwaegr.) Limpr.,
Laubm. Deutschl. 1: 600 (1888); Sim, Bryo. S. Afr.
241 (1926). Type: North America.
Barbula afrocaespitosa C. Mull, in Hedwigia 38:
109 (1899). Tortella afrocaespitosa (C. Mull.) Broth,
in Natiirl. PflFam. 1 : 397 (1902). Type: Cape, Kraka-
kamma, Ecklon s.n., 1832.
Barbula natalensicaespitosa C. Miill. in Hedwigia
38: 110 (1899); Tortella natalensicaespitosa (C. Miill.)
Broth, in Natiirl. PflFam. 1: 397 (1902). Type:
Natal, Gueinzius s.n.
Barbula eutrichostoma C. Miill. in Hedwigia 38: 110
(1899); Tortella eutrichostomum (C. Miill.) Broth, in
Natiirl. PflFam. 1: 397 (1902). Type: Cape, Blanco,
Rehmann s.n., 1875 (PRE!).
Tortella petrieana Sim, Bryo. S. Afr. 242 (1926).
Type: Natal, Glynn Falls, Sim 10063 (PRE, holo.!).
Plants small to medium, loosely caespi-
tose, green to yellow-green; terricolous,
saxicolous or corticolous. Stems 2-20 mm
tall, little branched; in section round, central
strand present, cortical cells thin-walled,
smaller, incrassate toward margin. Leaves
incurved, contorted with inrolled margins
dry, widespreading wet; oblong to oblong-
lanceolate, 3-5 mm long; apex obtuse to
broadly acute, mucronate; base scarcely
differentiated to oval; margins plane to
erect at apex, entire. Costa short-excurrent,
ventral superficial cells quadrate, papillose,
dorsal superficial cells linear, smooth; in
section subround, ventrally flattened, guide
cells 4, ventral stereid band strong, 2-3 cells
thick, ventral surface cells similar to laminal
cells, papillose, dorsal stereid band strong,
4-5 cells thick, dorsal surface cells not
differentiated. Upper laminal cells quadrate
Fig. 75. — Tortella humilis (1-10): 1. habit, x 1 ; 2. habit, x 5; 3. stem in cross section, x 170; 4. leaf, x 30;
5. leaf in cross section, x200 ; 6. cells at leaf base (right side), x 170; 7. upper laminal cells (papillae partly
shown), x 640 ; 8. leaf apex, x 170; 9. perigonium, x 60; 10. part of peristome and spores, x40. T. xanthocarpa
(linear leaf facies, 11-18): 11. habit, x 1; 12. habit, x5; 13. stem in cross section, x 170; 14. leaves, x30; 15.
leaf in cross section, x245; 16. cells at upper base (left side), xl70; 17. leaf apex, xl70; 18. upper laminal
cells, x 640. T. xanthocarpa (typical facies, 19-28): 19. habit, x 1 ; 20. habit x5;21.1eaf, x 30; 22. leaf in cross
section, x245; 23. cells at upper leaf base, xl70; 24. upper laminal cells, x640; 25. leaf apex, xl70; 26.
operculate capsule, x5; 27. part of capsule mouth with peristome, X200; 28. spore, x700. (1-10, Jacot Guil-
larmod 6171 ; 11-18, Magill 4812; 19-28, Magill 5983).
POTTIACEAE
257
258
POTTIACEAE
to subquadrate, occasionally angular, thin-
walled, superficially bulging, papillae low,
mostly 4 over lumen; basal cells rectangular,
hyaline, smooth, extending up margin, con-
fined to lower \ of leaf.
Autoicous. Perigonia gemmate, on short
lateral branches; perichaetia terminal, leaves
not differentiated. Seta 7-12 mm long,
yellowish; capsule cylindrical, 2-3 mm long,
yellow-brown; peristome to 1 mm high, red-
yellow, teeth filiform above very short basal
membrane, twisted 2-3 turns counterclock-
wise, papillose; operculum rostrate; calyptra
cucullate, covering upper \ of capsule;
spores round, 10-12 pm, papillose. Fig. 75;
1-10.
Tortella humilis is a very widespread species
found throughout Africa, America and parts of
Europe and Asia. In Southern Africa it is frequently
collected in the southwestern, southern and eastern
Cape, Transkei, Natal, Zululand, Lesotho, eastern,
central and northern Transvaal and Swaziland. Map
100.
Vouchers: Bosman 1608; Boucher 326 9b; Gold-
blatt 2124 d\ Killick 4206; Magill 4563; Watson 1359.
The plant shows some variation in leaf length,
in the distance the basal leaf cells extend up the mar-
gins and the habitat; but the oblong leaves with obtuse
apices separate T. humilis from other Southern Afri-
can species.
3. Tortella fragilis {Hook. & Wils.)
Limpr., Laubm. Deutschl. 1: 606 (1888);
Saito in J. Hattori bot. Lab. 39: 445 (1975).
Type: North America, Drummond 127.
Didymodon fragilis Hook. & Wils. in Drumm.,
Musci Bor. Am. 127 (1828).
Plants large, forming dense tufts or
cushions, dark to yellow-green; terricolous or
saxicolous. Stems 30-40 mm high, branching
above, radiculose below, dark red; in section
round, central strand absent, cortical cells
lax, smaller toward margin, outer cortical
cells in 2-3 rows, strongly thickened, reddish.
Leaves incurved to appressed, weakly twisted
dry, erect-spreading wet, tips fragile; linear-
lanceolate, 5-6 mm long; bistratose above;
ventral surface flat; apex subulate, frequently
broken above; base ovate to oblong; mar-
gins plane, entire, bordered above by single
row of smooth, incrassate rectangular cells.
Costa strong, excurrent, ventral superficial
cells quadrate, papillose below, long- rec-
tangular, smooth above, dorsal superficial
cells narrowly rectangular, smooth through-
Fig. 76. — Tortella fragilis: 1. habit, xl; 2.
stem in cross section, xl70; 3. leaf, x30; 4. leaf in
proximal cross section, x220; 5. leaf in distal cross
section, x220; 6. laminal cells at mid-leaf showing
larger basal cells extending up left margin, xl70;
7. upper laminal cells (papillae partly shown), x640;
8. leaf apex, X 170. (1—8, Magill 4408).
POTTIACEAE
259
out; in section semicircular, guide cells 8-10,
occasionally with 1-2 cells above or below
main layer, ventral stereid band strong, to 5
cells thick, ventral surface cells similar to
laminal cells, papillose in lower leaf, smooth
above, dorsal stereid band strong, 6-8 cells
thick, surface cells not differentiated or 1-2
cells at insertion of lamina larger, incrassate,
papillose. Upper laminal cells rounded,
quadrate to short-rectangular, incrassate,
papillae low, massive, mostly 4, completely
covering cells in upper leaf; marginal cells in
1-3 rows, rectangular, incrassate, smooth;
basal cells rectangular, lax, hyaline, smooth,
extending up margin, confined to lower ^ or
ending below mid-leaf.
Sporophytes unknown. Fig. 76: 1-8.
Although T. fragilis occurs throughout the
Northern Hemisphere it has not been reported pre-
viously for the Southern Hemisphere. The Southern
African plants were collected above 2 700 m in the
mountains of eastern Lesotho. Map 99.
Vouchers: Magill 4341, 4408.
The long, rigid, fragile leaf tip, bistratose upper
leaf and large size of the plants distinguish T. fragilis
from other species of Tortella. The phytogeographic
importance of eastern Lesotho is again demonstrated
by the presence of this Northern Hemisphere species
at high elevations in the Drakensberg.
5. OXYSTEGUS
Oxystegus ( Limpr .) Hilp. in Beih. bot. Zbl. 50: 666 (1933); Saito in J. Hattori bot. Lab. 39:
436 (1975); Gangulee, Moss. E. India 653 (1972). Lectotype species: O. cylindricus (Brid.)
Hilp.
Plants small to medium, yellow-green; saxicolous or terricolous. Stems without central
strand. Leaves linear-lanceolate to ligulate, fragile; apex acute; margins plane, entire. Laminal
cells quadrate to short-rectangular, papillose; basal cells lax, smooth.
Sporophyte terminal; capsule cylindrical; peristome teeth filiform, erect, distant, without
basal membrane.
Thirteen species of Oxystegus are known, primarily from southern Asia, Africa and South America.
Oxystegus cylindricus occurs throughout the Northern Hemisphere. The genus is recognized by the absence of a
central strand of the stem, narrow, fragile leaves with plane margin, a costa with dorsal and ventral stereid
bands, and a peristome without a basal membrane.
Oxystegus cylindricus {Brid.) Hilp. in
Beih. bot. Zbl. 50: 620 (1933); Saito in J.
Hattori bot. Lab. 39: 437 (1975). Type:
Europe.
Weissia cylindrica Bruch ex Brid., Bryol. Univ. 1 :
806 (1827). Trichostomum cylindricum (Brid.) C. Miill.,
Syn. Muse. 1: 586 (1849), non Hedwig (1801); Sim,
Bryo. S. Afr. 245 (1926).
Barbula leptotortella C. Miill. in Hedwigia 38: 110
(1899). Type: Cape, Somerset East, Boschberg,
MacOwan s.n.
Plants small to medium, loosely caespi-
tose, yellow-green; terricolous or saxicolous.
Stems 5-10 mm high, occasionally branched
above; in section round, central strand
absent, cortical cells incrassate, smaller
toward margin; axillary hair long, to 20
cells, hyaline throughout. Leaves crowded
above, incurved, contorted to spirally twisted
dry, erect-spreading wet, frequently fragile
or broken; linear-lanceolate or narrowly
ligulate, 2, 5-4, 5 mm long; apex acute to
narrowly acuminate; margins plane, entire.
Costa percurrent to short-excurrent, ventral
superficial cells quadrate, papillose, dorsal
superficial cells narrowly rectangular, smooth ;
in section semicircular, guide cells 4-5,
ventral stereid band strong, 2-3 cells thick,
ventral surface cells small, similar to laminal
cells, dorsal stereid band 2-4 cells thick,
dorsal surface cells undifferentiated or sub-
stereids. Upper laminal cells quadrate to
short-rectangular, incrassate, papillae low,
blunt, 4-8 over lumen ; basal cells rectangular,
hyaline, smooth, thin-walled, junction with
laminal cells straight across leaf or occasion-
ally extending a short distance up margin.
Dioicous. Perichaetia terminal, leaves
short, base broader than vegetative leaves.
Seta 6-8 mm long, yellow; capsule cylin-
drical, 2 mm long, reddish yellow; peristome
260
POTTIACEAE
variable, erect or weakly twisted, teeth fili-
form, occasionally perforated or divided
above, papillose; operculum long-rostrate,
0,8 mm long; calyptra cucullate, 2 mm long;
spores round, 8-12 //m, granulate. Fig. 77:
1-10.
Oxystegus cylindricus is almost cosmopolitan in
distribution, absent only from Australasia, Australia,
New Zealand and Antarctica. In Southern Africa,
plants are uncommon. They occur in Natal and in
scattered localities in the eastern and southwestern
Cape Province. Map 98.
Vouchers: Magill 3802, 3980, 5605; Owen 17;
Sim 10067.
Although some plants of O. cylindricus approach
Trichostomum brachydontium in habit and leaf shape
(see note under that species), the presence or absence
of the central strand in the stem is reliable in separat-
ing similar specimens. Most specimens of O. cylindri-
cus have the typical narrow, often fragile leaves, and
a more open appearance with incurled, twisted leaves
when dry. In addition, O. cylindricus has a well
developed peristome, while in Southern African
specimens of T. brachydontium peristomes are rudi-
mentary or absent.
6. TRICHOSTOMUM
Trichostomum Bruch in Flora, Jena 12: 295 (1829); Saito in J. Hattori bot. Lab. 39: 431
(1975); Gangulee, Moss. E. India 675 (1972). Type species: T. brachydontium Bruch.
Plants short to tall, dark green to yellow-green; saxicolous or terricolous. Stems with
well-developed central strand. Leaves narrowly spathulate to elliptical; apex broad; margins
plane, entire. Laminal cells quadrate to subquadrate, densely papillose; basal cells lax,
smooth.
Sporophyte terminal; capsule short-cylindrical; peristome present, absent or rudimentary,
when developed with basal membrane.
The 106 species of Trichostomum are equally distributed throughout the world. The genus is recognized
by the presence of a central strand in the stem, broad leaves with plane margins, short-excurrent costa with
dorsal and ventral stereid bands, and a peristome, with erect teeth from a short basal membrane, which is
frequently rudimentary or absent.
Trichostomum brachydontium Bruch ex
F. A. Muell. in Flora, Jena 12: 393 (1829);
Saito in J. Hattori bot. Lab. 39: 431 (1975);
Smith, Moss FI. Brit. Irel. 291 (1978); Sim,
p.p., Bryo. S. Afr. 242 (1926). Type: Europe.
Pottia zeyheri Hampe ex C. Mull., Syn. Muse. 1 :
561 (1849). Hyophila zeyheri (Hampe) Jaeg. in Verh.
St Gall, naturw. Ges. 1871-72: 355 (1873); Sim,
Bryo. S. Afr. 221 (1926). Type: Cape, Zeyher s.n.
(BM!).
Pottia afrophaea C. Mull, in Hedwigia 38: 97
(1899); Sim, Bryo. S. Afr. 219 (1926). Hyophila
afrophaea (C. Mull.) Warnst. in Hedwigia 58: 59
(1916). Type: Orange Free State, Bethlehem, Reh-
mann 1 20 (PRE!).
Plants medium-sized, caespitose, dark
green to yellow-green; terricolous or saxi-
colous. Stems 4-26 mm tall ; in section round,
central strand present, frequently large,
cortical cells ± incrassate, smaller toward
margin, occasionally stereids; axillary hairs
8-12 cells long, hyaline throughout. Leaves
incurved, appressed to crisped dry, widely
spreading wet; narrowly spathulate to ellip-
tical or oblong, 2, 5-3, 5 mm long; apex
broadly acute to obtuse, cuspidate; margins
plane, entire. Costa short-excurrent, ventral
superficial cells quadrate, papillose, dorsal
superficial cells rectangular, smooth; in
section semicircular to subround, guide cells
4, ventral stereid band strong, 2-3 cells
thick, ventral surface cells similar to laminal
cells, dorsal stereid band strong, 3-4 cells
thick, dorsal surface cells substereids or
differentiated only at insertion of lamina.
Fig. 77. — Oxystegius cylindricus (1-10): 1. habit, X 1 ; 2. habit, x8; 3. stem in cross section, Xl70; 4.
leaf, x40; 5. leaf in cross section, X 130; 6. cells at leaf base (right side), X 170; 7. upper laminal cells (papillae
partly shown), x 640; 8. leaf apex, x 170; 9-10. parts of capsule mouth with peristome teeth, X 170. Trichosto-
mum brachydontium (1 1-19): 11 12. habit, xl; 13. habit, x 8; 14. stem in cross section, x 170; 15. leaves, x40;
16. leaf in cross section, x 130; 17. cells at leaf base (right side), x 170; 18. upper laminal cells (papillae partly
shown), X 640; 19. leaf apex (dorsal surface), X 170. (1-10, Magill 3980; 1 1 & 13-19, Sim 8579; 12, Smook 944).
POTTIACEAE
261
262
POTTIACEAE
Upper laminal cells quadrate to subquadrate,
incrassate, papillae numerous, low, blunt,
scattered over lumen; marginal cells generally
papillose; basal cells rectangular, hyaline,
smooth, thin- walled, junction with laminal
cells ± straight across leaf.
Dioicous. Perichaetia terminal, leaves
undifferentiated. Seta 5-10 mm long, yellow-
Map 101. — • Trichostomum brachydontium
ish; capsule short-cylindrical, 1,5 mm long;
peristome absent or rudimentary; operculum
rostrate, 1 mm long; calyptra cucullate;
spores round, 15-17 pm, papillose. Fig. 77:
11-19.
A cosmopolitan species, T. brachydontium is
also widespread in Southern Africa and found in a
large variety of habitats. Map 101.
Vouchers: Crosby & Crosby 7699; Hardy 5192;
Lavranos 15204a; Magill 4545a, 5447, 6048; Rankin
12; Retief 360a; Vahrmeijer PRE-CHI 2674; Van
Rooy 56, 269.
A degree of variability is expressed by T. brachy-
dontium in Southern Africa. The specimens generally
reflect differences in habitat through stature of the
plants. The leaves also vary in size and to some
degree shape, but plants from desert or forest are
recognized by the narrowly spathulate or elliptical
leaves with broad apices. In Southern African speci-
mens the peristome is absent or rudimentary, a con-
dition also known from South American specimens.
Northern Hemisphere specimens generally have
short, fragile teeth.
Insufficiently Known Species
Trichostomum leiodontium C. Mull, in Hedwigia
38: 100 (1899). Type: Cape, Somerset East, Bosch-
berg, MacOwan s.n. The type has not been seen.
Brotherus in Natiirl. PflFam. 1: 397 (1902) con-
sidered this species a synonym of Tortella xantho-
carpa (C. Mull.) Broth.
7. WEISSIA
Weissia Hedw., Spec. Muse. 64 (1801); Saito in J. Hattori bot. Lab. 39: 417 (1975); Smith,
Moss FI. Brit. Irel. 274 (1978). Type species: W. controversa Hedw.
Hymenostomwn R. Br. in Trans. Linn. Soc. Lond. 12: 572 (1819); Sim, Bryo. S. Afr. 254 (1926). Type
species: H. microstomum (Hedw.) R. Br.
Plants small, in loose tufts, green to yellow-green; terricolous. Stems short; central strand
present. Leaves larger above, incurved-contorted or twisted above base when dry, spreading
wet, narrow; linear, lanceolate or oblong; apex acute to obtuse, frequently cucullate, mucro-
nate to apiculate; margins involute to incurved. Costa short-excurrent, rarely percurrent in
some leaves, ventral superficial cells quadrate, papillose, frequently patchy, rarely absent; in
section with dorsal and ventral stereid bands. Laminal cells quadrate, weakly thickened, papil-
lose, occasionally papillae on dorsal and ventral surfaces differing.
Autoicous or paroicous (? dioicous). Perichaetial leaves little differentiated, somewhat
longer. Seta erect, yellowish; capsule cylindrical; peristome present, rudimentary or absent,
capsule occasionally hymenostomaceous; operculum rostrate, cells not twisted; calyptra
cucullate; spores round, papillose.
The genus Weissia, as circumscribed here, comprises the species included by some authors in Hymenosto-
mum. The division lines between capsules that are either hymenostomaceous or gymnostomaceous and those
producing a rudimentary or complete peristome is not altogether clear, and reliable gametophytic characters
are not always available. Therefore all species present in Southern Africa are treated under Weissia, and the
genus is divided into the subgenera Weissia and Hymenostomum.
POTTIACEAE
263
The genus Weissia consists of c. 88 species found throughout the world. A few species, such as W. contro-
versa, are widespread to cosmopolitan, but the great majority of species is very restricted or even local in distri-
bution. This is the case with the Southern African species. A broader revision and more realistic interpretation
of sporophyte and peristome development will no doubt greatly reduce the number of species presently recog-
nized in the genus.
1 Peristome present, occasionally very rudimentary (subgen. Weissia ):
2 Plants paroicous; peristome teeth linear, distant; leaves to 2 mm long; costa percurrent to very short-
excurrent 1. W. dieterlenii
2 Plants autoicous; peristome teeth lanceolate, frequently irregularly cleft or perforated; leaves to 4 mm
long; costa short-excurrent 2. W. controversa
1 Peristome absent (subgen. Hymenostomum ):
3 Leaves linear-lanceolate, 3-4 mm long; costa apiculate 3. W. humicola
3 Leaves to 2 mm long; costa percurrent to mucronate:
4 Leaves narrowly lanceolate; ventral superficial costal cells elongate, smooth throughout. .4. W. cucullata
4 Leaves oblong to narrowly oval; ventral superficial costal cells quadrate, papillose 5. W. latiuscula
1. Weissia dieterlenii Ther. in Bull.
Mus. Hist, nat., Paris 30: 240 (1924); Broth,
in Natiirl. PflFam. 10: 255 (1924). Type:
Lesotho, Leribe, Dieterlen s.n. (PC, holo.!;
PRE!; NH!).
Weissia brachycarpa C. Mull, in Hedwigia 38: 112
(1899), horn, illeg., non (Nees & Hornsch.) Jur. (1882).
Hymenostomum brachycarpum (C. Miill.) Par., Ind.
Bryol. Suppl. 189 (1900); Broth, in Natiirl. PflFam.
10: 254 (1924). Weissia viridula Hedw. var. brachy-
carpa (C. Mull.) Dix. in Trans. R. Soc. S. Afr. 18:
251 (1929) [excluding Wager 988; Pillans 4748],
non Weissia viridula Hedw. var. brachycarpa Nees &
Hornsch. (1831). Weissia controversa var. pillansii
Schelpe in Trans. R. Soc. S. Afr. 44: 116 (1979).
Type: Orange Free State, Renoster River, Rehmann
s.n., Herb. Jack (G. holo.!).
Plants small, loosely caespitose, yellow-
green; terricolous. Stems 2-4 mm tall; in
section round, central strand large, inner
cortical cells large, in single row, outer corti-
cal cells in 2-3 rows, smaller. Leaves spirally
twisted dry, erect-spreading wet; lanceolate,
1, 5-2,0 mm long; apex acute, mucronate,
frequently obtuse in lower leaves; base oval
to oblong; margins narrowly involute, entire.
Costa percurrent to very short-excurrent,
ventral superficial cells quadrate, papillose,
dorsal superficial cells rectangular, smooth;
in section round, guide cells 2, ventral stereid
band of 1-2 cells only, ventral surface cells
similar to laminal cells, dorsal stereid band
2-3 cells thick, occasionally substereids.
Upper laminal cells quadrate to subquadrate
or angular, weakly incrassate, superficially
bulging on both surfaces, papillae low and
simple; basal cells rectangular, hyaline,
smooth, thin-walled.
Paroicous. Antheridia in upper leaf
axils; perichaetia terminal, leaves undifferen-
tiated. Seta 6-7 mm long, yellow; capsule
short-cylindrical, 1,2 mm long, reddish
yellow; peristome present, inserted below
mouth, teeth linear, distant, 120 pm high,
erect, red-yellow, weakly papillose; oper-
culum rostrate, 0,8 mm long; spores sub-
round to angular, (12-) 15-17 pm, irregularly
papillose. Fig. 78: 1-12.
Endemic to Southern Africa, W. dieterlenii is
known from grasslands of Lesotho and the Orange
Free State. Map 102.
Map 102. — • Weissia dieterlenii
x Weissia humicola
Voucher: Magill 4679; Van Rooy 366, 432.
Specimens of W. dieterlenii bear some resem-
blance to W. controversa, but the paroicous condition
and distinctive peristome will easily separate the two
species. See note on W. brachycarpa under W.
humicola.
264
POTTIACEAE
POTTIACEAE
265
2. Weissia controversa Hedw., Spec.
Muse. 67 (1801); Saito in J. Hattori bot. Lab.
39: 426 (1975); Smith, Moss FI. Brit. Irel.
276 (1978). Type: Europe.
Weissia viridula Hedw. ex Brid., hom. illeg., Bryol.
Univ. 1 : 364 (1826); Broth, in Natiirl. PflFam. 10: 255
(1924); Sim, Bryo. S. Afr. 253 (1926).
? Weissia linguaelata Shaw in Cape Monthly Mag.
17: 378 (1878). Type: Cape, Graaff-Reinet, McLea
s.n.
Weissia vallis-gratiae C. Mull, in Hedwigia 38: 111
(1899); Broth, in Natiirl. PflFam. 10: 255 (1924).
Type: Cape, Gnadenthal, Breutel s.n. Herb. Hampe
(BM, holo.!).
Weissia oranica C. Mull, in Hedwigia 33: 112
(1899); Broth, in Natiirl. PflFam. 10: 255 (1924).
Type: Orange Free State, ‘Blumfontein’, Rehmann 19
(BM!; PRE!).
Weissia viridula Hedw. var. longifolia Broth. &
Wag. in Trans. R. Soc. S. Afr. 4: 6 (1914); Sim,
Bryo. S. Afr. 253 (1926).
Plants small, loosely caespitose, green to
yellow-green; terricolous. Stems 2-4 mm
high, infrequently branched below; in section
round, central strand present, occasionally
large, inner cortical cells large, slightly
thickened, outer cells smaller. Leaves twisted
to spirally involute dry, widely spreading wet;
lanceolate to linear-lanceolate, 2-4(-5) mm
long; apex acute, apiculate; base ovate to
oblong; margins narrowly involute, entire.
Costa short-excurrent, ventral superficial
cells quadrate, papillose, frequently patchy,
exposing smooth, rectangular cells, dorsal
superficial cells long-rectangular, smooth; in
section round, guide cells 4, ventral stereid
band 1-2 cells thick, ventral surface cells
large, papillose, dorsal stereid band 2-3 cells
thick, dorsal surface cells undifferentiated,
smooth. Upper laminal cells quadrate to
rounded, thickened, papillae low, numerous,
scattered over lumen, dorsal surface occasion-
ally strongly thickened, obscuring papillae;
basal cells rectangular, hyaline, smooth.
Autoicous. Perigonia on short lateral
branches; perichaetia terminal, leaves similar
to vegetative leaves, frequently longer. Seta
(4-) 6-8 mm long, yellow; capsule short-
cylindrical, 1,5 mm long, reddish yellow,
mouth red; peristome short, inserted below
mouth, teeth lanceolate, obtuse, frequently
cleft or perforated, occasionally rudimentary,
150-200 pm high, reddish, papillose; oper-
culum rostrate, 0,5-0, 6 mm long; spores
round, 20-22 pm, strongly papillose. Fig.
78: 13-23.
Weissia controversa is a cosmopolitan species
frequently collected in eastern and southern parts of
Southern Africa. The small moss is common in city
gardens as well as natural habitats. Map 103.
Vouchers: Cholnoky 792; Esterhuysen 15532;
Eyles 1412; Magill 5871; Schelpe 7843; Wager 1447.
The long, narrow leaves with strongly involute
margins distinguish W. controversa from most other
mosses. The short, somewhat fragile peristome may
appear absent in older capsules but careful observa-
tion generally reveals basal fragments of the peri-
stome within the capsule mouth. Weissia humicola
and W. dieterlenii are both gametophytically similar
to W. controversa and identification is difficult with-
out capsules.
Fig. 78. — Weissia dieterlenii (1-12): 1. habit, xl; 2. habit, x20; 3. stem in cross section, x340; 4-5.
leaves, x40; 6. leaf in cross section, x 640; 7. cells at leaf base (left side), x 170; 8. upper laminal cells (papillae
partly shown), x640; 9. leaf apex, xl70; 10. perichaetial leaf with attached antheridium, x40; 11. part of
capsule mouth with peristome teeth and spores, X 170; 12. spore, x 640. W. controversa (13-23): 13. habit, x 1 ;
14. habit, x20; 15-16. leaves, x40; 17. leaf in cross section, x640; 18. cells at leaf base (right side), xl70;
19. upper lamina cells (papillae partly shown), >.640; 20. leaf apex, x 170; 21. perigonium, x75; 22-23. parts
of capsule mouth with peristome teeth, x 170. W. humicola (24—32) : 24. habit, x 1 ; 25. habit, x 10; 26. leaves,
x 40; 27. leaf in cross section, X 640; 28. cells at leaf base (right side), X 170; 29. upper laminal cells (papillae
partly shown), X 640; 30. leaf apex, X 170; 31. part of capsule mouth, x 170; 32. spore, X 640. (1-12, Dieterlen
s n.; 13-22, Sim 9180; 23, Eyles 1412; 24-32,/. Sim PRE-CH7846).
266
POTTIACEAE
3. Weissia humicola C. Mull, in Hed-
wigia 38: 112 (1899). Type: Cape, Somerset
East, Boschberg, MacOwan s.n., 1876.
Hymenostomum humicola (C. Mull.) Par., Ind.
Bryol. Sappl. 189 (1900); Broth, in Natiirl. PflFam.
10: 254 (1924); Sim, Bryo. S. Afr. 254 (1926).
Weissia viridula Hedw. var. brachycarpa sensu
Dixon in Trans R. Soc. S. Afr. 18: 251 (1929).
Plants small, loosely caespitose, yellow-
green; terricolous. Stems 2-5 mm tall,
branching below; in section round, central
strand present, cortical cells variable in size,
incrassate. Leaves contorted to incurved dry,
widely spreading wet; linear-lanceolate, 3-4
mm long; apex acute, apiculate; base
oblong; margins narrowly incurved to invo-
lute, entire. Costa short-excurrent, ventral
superficial cells quadrate, papillose, occasion-
ally patchy, exposing smooth, rectangular
cells, dorsal superficial cells long-rectangular,
smooth; in section subround, guide cells 2-4,
ventral stereid band weak, of 1-2 cells only,
ventral surface cells similar to laminal cells,
papillose or stereids exposed, smooth, dorsal
stereid band 2-3 cells thick, dorsal surface
cells undifferentiated. Upper laminal cells
quadrate, thickened, papillae low, numer-
ous, scattered over lumen; basal cells rec-
tangular, hyaline, smooth, thin-walled.
?Dioicous. Perigonia not seen; peri-
chaetia terminal, leaves similar to vegetative
leaves, but slightly longer. Seta 4-5 mm long,
yellow; capsule short-cylindrical, 1 mm long,
yellowish; peristome absent; operculum ros-
trate, 0,5-0, 6 mm long; spores round,
17-20 pm, strongly papillose. Fig. 78: 24-32.
Endemic to Southern Africa, W. humicola is
known only from shrublands of the eastern Cape
Province. Map 102.
Vouchers: MacOwan 22; J. Sim PRE-CH7846.
Specimens of W. humicola are very similar to W.
controversa and may represent a gymnostomous form
of that species, since the capsules examined were not
hymenostomaceous. The present taxonomic trend is
to separate such taxa into different species and this is
followed here. Both species generally produce nume-
rous sporophytes, so that identification is possible.
Specimens of W. controversa with rudimentary
peristomes could easily be mistaken for this species.
The specimens ( Wager 988 ; Pillans 4748) cited by
Dixon (1929) when he made the combination W.
viridula var. brachycarpa (C. Mull.) Dix. are, on the
basis of gymnostomous capsules, best placed here.
The type on which the name is based (Rehmann s.n..
Herb. Jack, G!) has a distinctive peristome and has
been referred to W. dieterlenii Thdr.
4. Weissia cucullata C. Mull, in Bot. Ztg 16:
163 (1958). Type: Cape, Gnadenthal, Breutel
s.n., Herb. Hampe (BM, holo.l).
Hymenostomum cucullatum (C. Mull.) Kindb.,
Enum. Bryin. Exot. 62 (1888); Broth, in Natiirl.
PflFam. 10: 254 (1924).
Plants small, loosely caespitose, yellow-
green; terricolous. Stems 2-4 mm tall,
occasionally branched below; in section
round, central strand small, cortical cells
incrassate, outer row smaller. Leaves con-
torted-incurved dry, erect-spreading wet;
linear-lanceolate, to 2 mm long; apex acute,
weakly cucullate, mucronate; base oblong;
margins narrowly incurved, entire. Costa
short-excurrent, ventral superficial cells elon-
gate, smooth, dorsal superficial cells long-
rectangular, smooth; in section oval, guide
cells 4, ventral stereid band strong, 2-3 cells
thick, ventral surface cells undifferentiated,
dorsal stereid band 3-4 cells thick, dorsal
surface cells undifferentiated. Upper laminal
cells quadrate, thin-walled, papillae on
ventral surface large, branched, 1-2 over
lumen, on dorsal surface low, numerous,
scattered; basal cells rectangular, hyaline,
smooth, thin-walled.
Autoicous. Perichaetia terminal, leaves
similar to vegetative leaves, but slightly
larger. Seta 2-3 mm long, yellow; capsule
hymenostomaceous, oval, constricted at
mouth, 0, 8-1,0 mm long, reddish yellow;
peristome absent; operculum rostrate, 0,3-
Map 104. — • Weissia latiuscula
x Weissia cucullata
POTTIACEAE
267
0,4 mm long; spores round, 25 pm, strongly
papillose. Fig. 79: 1-9.
Endemic to Southern Africa, this species is
known only from the type specimen. The original
collection was made by Breutel, in the mountains
near the mission station at Gnadenthal, in the south-
western Cape. Map 104.
Voucher: Type only.
The small, linear-lanceolate leaves, acute-
cucullate apices and hymenostomaceous capsules
help to identify W. cucullata. The spores are also
larger than those of any other Southern African
species of Weissia.
5. Weissia latiuscula C. Mull, in Hedwigia
38: 111 (1899). Type: Transvaal, Lydenburg,
W ilms s.n., 1887 (G, holo. !).
Hymenostomum latiusculum (C. Miill.) Par., Ind.
Bryol. Suppl. 189 (1900); Broth, in Natiirl. PflFam.
10: 254 (1924).
Hymenostomum eurybasis Dix. in S. Afr. J. Sci.
18: 333 (1922); Sim, Bryo. S. Afr. 255 (1926). Syn-
types: Zimbabwe, Matopos, Eyles 940, 941 ; Magude,
Sim 8989.
Plants small, loosely caespitose, light
green to yellow-green; terricolous or saxi-
colous. Stems 2-4 mm high, branching below;
in section round, central strand large or
small, cortical cells lax, outer 1-2 rows
smaller. Leaves closely incurved dry, widely
spreading wet; oblong to narrowly oval,
1 , 8-2 , 5 mm long, upper and subperichaetial
leaves linear; apex rounded-obtuse, cucullate,
mucronate; base oblong; margins broadly
incurved, entire. Costa percurrent to short-
excurrent, ventral superficial cells quadrate,
papillose, dorsal superficial cells rectangular,
smooth; in section round, guide cells 2-4,
ventral stereid band 2-3 cells thick, ven-
tral surface cells similar to laminal cells,
papillose, dorsal stereid band 4 cells thick,
dorsal surface cells undifferentiated, smooth.
Upper laminal cells quadrate to subquadrate,
ventral surface bulging, papillae numerous,
Fig. 79. — Weissia cucullata (1-9): 1. habit, xl;
2. habit, x 10; 3. stem in cross section, x275; 4.
leaves, x40; 5. leaf in cross section, x530; 6. cells at
leaf base, xl70; 7. upper laminal cells (papillae
partly shown), x640 ; 8. leaf apex, xl70; 9. spore,
x640. W. latiuscula (10-17): 10. habit, xl; 11. habit,
X10; 12-13. leaves, x40; 14. leaf in cross section,
x340; 15. basal leaf cells at right margin, xl70;
16. upper laminal cells (papillae partly shown), x 640;
17. leaf apex, xl70. (1-9, Breutel s.n.; 10-17, Smook
& Harding 705a).
268
POTTIACEAE
low, simple or raised on incrassate mam-
millae, dorsal surface flat, papillae numerous,
low, simple; basal cells rectangular, hyaline,
smooth, thin-walled.
Autoicous. Perigonia on short lateral
branches; perichaetia terminal, leaves linear,
acute. Seta to 4 mm long, yellow; capsule
cylindrical, 1 mm long, conspicuously con-
stricted below mouth dry, yellowish, mouth
red; peristome absent; operculum rostrate,
0,6 mm long; spores round, 20-22 /im,
papillose. Fig. 79: 10-17.
In Southern Africa W. latiuscula is known from
dry shrublands of central South West Africa/Nami-
bia, the northern, central and southeastern Transvaal
and northern and central Cape. The species has also
been collected in Zimbabwe. Map 104.
Vouchers: Smook 72a, 705a; Wager 345.
The oblong leaves with obtuse apices and broadly
incurved margins distinguish this species from other
species of Weissia in Southern Africa. The plants
generally show considerable variation between sterile
and fertile plants. This applies particularly to leaf
size and shape. The leaves of sterile plants and
lower leaves of fertile plants are oblong-obtuse, while
subperichaetial and perichaetial leaves are linear
with acute apices.
269
BR Y OB ARTR AMI ACE AE
Plants small on persistent protonema, in small groups; terricolous. Stems erect; central
strand absent. Leaves larger above, narrow, ligulate to Ungulate; apex obtuse in lower leaves
to subulate in subperichaetial leaves; margins plane, papillose. Costa ending below apex or
ending in subula of subperichaetial leaves; in section with dorsal stereid or substereid band.
Upper laminal cells quadrate to subhexagonal, papillose; basal cells rectangular, smooth.
Paroicous. Perichaetia terminal, leaves linear-lanceolate, subulate. Seta short, erect;
capsule cleistocarpic, urn globose-apiculate; epigonium persistent; spores large, warty.
BRYOBARTRAMIA
Bryobartramia Sainsb. in Bryologist 51 : 10 (1948); Scott & Stone, Moss. S. Aust. 220 (1976);
Stone in Aust. J. Bot. 25: 141 (1977). Type species: B. robbinsii Sainsb.
With characters of the family.
Bryobartramia is a monotypic genus presently known only from Australia and Southern Africa. The genu s
shows similarities to genera of Pottiaceae and Encalyptaceae, but is recognized as a separate family because of
its persistent epigonium. In all other species of mosses the epigonium splits into a lower part, the vaginula, that
surrounds the lower seta and foot of the sporophyte and an upper part, the calyptra, that covers the developing
capsule. As the seta elongates the calyptra is pushed away from the vaginula. In Bryobartramia the epigonium
enlarges to accommodate the entire sporophyte without splitting. Release of spores occurs only after the capsule
and epigonium deteriorate. The epigonium is known to frequently outlast the capsule.
Bryobartramia novae-valesiae {Broth.)
Stone & Scott in J. Bryol. 7: 604 (1973);
Stone in Aust. J. Bot. 25: 141 (1977). Type:
Australia, New South Wales, Cowra, Watts
7829 (NSW).
Trachycarpidium novae-valesiae Broth, ex Roth in
Hedwigia 53: 94 (1913).
Bryobartramia robbinsii Sainsb. in Bryologist 51:
10 (1948); Stone & Schelpe in J1 S. Afr. Bot. 39: 131
(1973). Type: Australia, Victoria, Castlemaine,
Robbins s.n., 1941.
Plants small, scattered or in small
groups, light green to yellowish with age;
terricolous. Stems to 3 mm tall, occasionally
branched above; in section round, central
strand absent, cortical cells thin-walled, in 4
rows, not differentiated. Leaves somewhat
flexuose above base dry, erect-spreading wet,
narrow; lower leaves variable in shape,
ligulate to lingulate, 0,3-0, 5 mm long;
apex obtuse; margins plane, entire or some-
what irregular, costa subpercurrent ; upper
leaves linear, ligulate, lingulate or lanceolate,
1,0-1, 2 mm long; apex broadly acute to
obtuse; margins plane, papillose. Costa
percurrent or ending below apex, weak in
leaf base, ventral superficial cells quadrate,
papillose, similar to laminal cells, dorsal
superficial cells long-rectangular, smooth; in
section subround, guide cells 2, incrassate,
ventral cells large, in single row, papillose,
dorsal stereid or substereid band 2-3 cells
thick, frequently exposed, dorsal surface
cells large, incrassate, papillose. Upper laminal
cells quadrate to subhexagonal, weakly
thickened, with 2-4 C-shaped papillae over
lumen; basal cells rectangular, thin-walled,
smooth.
Paroicous. Perichaetia terminal on stem
or branches; leaves linear-lanceolate, 1,8-
2,0 mm long; apex subulate; margins plane,
entire or sparsely denticulate in subula;
costa filling subula. Seta 0,2-0, 4 mm long,
red-brown; capsule cleistocarpic, urn globose
to oval, apiculate, 0,8 mm long, yellowish;
exothecial cells isodiametric, thin-walled,
stomata phaneropore, scattered over urn;
epigonium persistent, oval-rostrate, 1,7-2, 2
mm long, cells below beak prorate; spores
subround, 35-40 pm, warty, yellow-brown.
Fig. 80: 1-13.
Bryobartramiaceae
Map 105. — • Grimmia apocarpa
X Bryobartramia novae-valesiae
This species is known from southern Australia
and Southern Africa. In the Flora area, B. novae-
valesiae has been collected only a few times in the
western Cape Province. The specimens were collected
on clay banks in dwarf-succulent shrublands of the
Cedarberg and Richtersveld, and in the Karoo at
Worcester. Map 105.
Vouchers: Magill & Schelpe 3850; Oliver et al.
PRE-CH12887; Schelpe 7622; Van Zanten et al.
7608306d.
Specimens may be confused macroscopically
with Goniomitrium, but the narrow, papillose leaves
and persistent epigonium distinguish this species.
Fig. 80. — Bryobartramia novae-valesiae: 1. habit,
xl; 2. habit, x5; 3. stem in cross section, xl80;
4-5. leaves, x40; 6. leaf in cross section, x350; 7.
cells at leaf base (right side), x 170; 8. upper laminal
cells, x350; 9. leaf apex, dorsal surface (papillae
partly shown), xl70; 10. axillary hair, x200; 11.
perichaetial leaf, X40; 12. epigonium enclosing
capsule, x20; 13. spore, X640. (1-13, Schelpe 7672).
271
GRIMMIACEAE
Plants small to large, forming dense turfs or cushions, blackish green to yellow-green;
mostly saxicolous. Stems erect; central strand present or absent. Leaves ovate to lanceolate;
apex acute to acuminate, often with hyaline tips or awns; lamina frequently bistratose or only
margins thickened. Costa percurrent to excurrent; in section cells frequently undifferentiated.
Laminal cells quadrate to rectangular, incrassate, smooth, lateral walls frequently sinuate to
nodose; basal cells elongate, walls sinuate to straight.
Perichaetia terminal. Seta short, erect or cygneous ; capsule immersed or exserted, ovoid
to cylindrical; peristome single, teeth 16, cleft or perforated, reddish; operculum rostrate;
calyptra mitriform; spores small.
1 Lateral walls of basal cells sinuate to nodose ; stems without central strand. . . 3. Racomitrium
1 Lateral walls of basal cells smooth to weakly sinuolate; stems with central strand:
2 Plants small, blackish; perichaetia conspicuous, hyaline, perichaetial leaves strongly
differentiated; capsule immersed, annulus persistent 1. Leucoperichaetium
2 Plants small to medium, dark green to yellow-green; perichaetia not distinct; capsules
exserted or immersed, annulus deciduous 2. Grimmia
1. LEUCOPERICHAETIUM
Leucoperichaetium Magill, gen. nov., ad Grimmiam accedens sed ab eo cellulis laminae parietibus
plerumque rectis, foliis perichaetii distinctissimis et annulo persistenti super ostium thecae ubi
sicco inflexo differt.
Plantae parvae pro parte maxima infossae, tapetiformes, nigrae ad atrovirides; saxicolae
vel terricolae. Rami in parte superiore ramificantes ; filo medio. Folia parva, mutica; lamina
bistratosa supra basim. Costa percurrens, grege substereidarum dorsalium. Celluli laminae
quadrati, parietibus plerumque rectis.
Perichaetia terminalia conspicua; foliis distinctissimis, piliferis, hyalinis. Seta brevissima;
theca immersa cupulata; peristomio rudimentali papilloso; annulo persistanti, erect o ubi humido
inflexo super ostium ubi sicco; operculo convexo-apiculato ; calyptra parva mitriformi.
Type species: L. eremophilum Magill.
Plants small, mostly buried, forming turfs, blackish to blackish green; saxicolous or
terricolous. Stems branching above; central strand present. Leaves small, muticous; lamina
bistratose above base. Costa percurrent, with dorsal substereid band. Laminal cells quadrate,
walls mostly straight.
Perichaetia terminal, conspicuous; leaves strongly differentiated, piliferous, hyaline.
Seta very short; capsule immersed, cupulate; peristome rudimentary, papillose; annulus
persistent, erect wet, indexed over mouth dry; operculum convex-apiculate; calyptra small,
mitriform.
Similar to Grimmia but differing in mostly straight-walled laminal cells, strongly differentiated perichaetial
leaves and persistent annulus that inflexes over capsule mouth when dry.
Ill
Grimmiaceae
Grimmiaceae
273
Leucoperichaetium eremophilum Magill,
sp. nov., bene distincta dignoscenda plantis
parvis pro parte maxima infossis, nigribus;
perichaetiis magnis hyalinis; foliis vegetativis
oblongo-acuminatis, 0,8- 1,2 mm longis, mu-
ticis, bistratosis supra basim ; foliis perichae-
tiorum late ovalibus, abrupte piliferis, hyalinis;
theca immersa, peristomio rudimentali ; annulo
persistenti, alto cellulis quattuor, hygroscopico.
Type: South West Africa/Namibia, Wit-
piitz, on quartzite outcrop, Hardy 4864
(PRE, holo.; H; MO; NY; US).
Plants small, in irregular turfs, blackish
green; saxicolous. Stems erect, 5-10 mm long,
mostly buried, frequently branching; in
section round, central strand large, inner
cortical cells in 3-4 rows, thin-walled, outer
row slightly smaller, weakly thickened.
Leaves weakly appressed to incurved dry,
patent wet, crowded above; oblong-acumi-
nate, 0,8- 1,2 mm long, gradually larger
above on fertile plants, subperichaetial leaves
to 1,5 mm long; apex acute; margins erect,
entire; lamina in distal section bistratose, in
proximal section unistratose or with small
bistratose areas. Costa percurrent to just
excurrent as weak hyaline tip; in proximal
section oval, guide cells 4, slightly thickened,
ventral cells in single row, similar to laminal
cells, dorsal substereid band weak, 1-2 cells
thick, dorsal surface cells similar to laminal
cells, outer walls strongly thickened, in distal
section oval, guide cells 2, dorsal substereid
band present, reduced to 4-6 cells, dorsal and
ventral surface cells similar to laminal cells,
incrassate. Upper laminal cells rounded-
quadrate, incrassate, smooth; basal cells
larger, quadrate to rectangular, thin-walled.
Cladautoicous. Perigonia terminal;
leaves broadly ovate-acuminate, 0 , 6-0 , 7 mm
long; perichaetia terminal, strongly differen-
tiated, conspicuous ; leaves gradually modified
toward apex, outer perichaetial leaves dark
green with hyaline bases; obovate, abruptly
acuminate; margins entire; upper laminal
cells rounded quadrate, incrassate; basal
cells larger, rectangular, thin-walled, extend-
ing up margins to above mid-leaf; inner
perichaetial leaves light green to hyaline
throughout; broadly oval, abruptly pili-
ferous; margins irregularly serrate at apex;
costa short-excurrent; upper laminal cells
rhombic, incrassate; basal cells larger, rec-
tangular, very thin-walled, occupying lower f
of leaf. Seta erect, 0,3 mm long; capsule
immersed, urn cupulate, 0,7 mm long, mouth
wide; exothecial cells rectangular to oblong-
hexagonal, thin-walled, stomata large,
restricted to base of capsule; annulus per-
sistent, cells subhexagonal, very strongly
thickened, to 4 cells high, inflexed and nar-
rowing mouth when dry, upright when wet;
peristome rudimentary, teeth irregular, just
visible above annulus, ornately papillose;
Map 106. — • Grimmia ovalis
x Leucoperichaetium eremophilum
Fig. 81. — Leucoperichaetium eremophilum (1-18): 1. habit, xl; 2. habit (soil level shown), x5; 3. vege-
tative leaves, x40; 4. upper leaves from fertile branch, x40; 5. subperichaetial leaf, x40; 6. perichaetial leaf,
x40; 7. leaf in proximal cross section, x240 ; 8. leaf in lower median cross section, x240; 9. leaf in upper
median cross section, x 240; 10. leaf in distal cross section, x 240; 11. cells at leaf base (left side), x 170; 12.
cells at perichaetial leaf base (right side), x 170; 13. apex of vegetative leaf, x 170; 14. apex of upper leaf from
fertile branch, xl70; 15. apex of subperichaetial leaf, xl70; 16. apex of perichaetial leaf, xl70; 17. capsule
and perichaetial leaf, xl5; 18. part of capsule mouth with persistent annulus and peristome teeth, x310.
Grimmia apocarpa (19-29): 19. habit, wet, xl; 20. habit, dry, xl; 21. habit, x5; 22. muticous leaves, x20;
23. piliferous leaf, x20; 24. leaf in cross section, xl30; 25. cells at leaf base (right side), xl30; 26. upper
laminal cells at right margin, x 130; 27. leaf apex, x 130; 28. part of capsule mouth with peristome teeth, x 240;
29. capsule with perichaetial leaves, x 10. (1-8, Hardy 4864; 19-22 & 24-29, Magill 4197; 23, Liebenberg 5848).
274
Grimmiaceae
operculum low-convex, bluntly apiculate;
calyptra mitrate, smooth, covering upper
capsule; spores subround, 12-15 pm, granu-
late. Fig. 81: 1-18.
Endemic to Southern Africa, L. eremophilum is
presently known only from quartzite outcrops in the
dwarf succulent shrublands of southern South West
Africa/Namibia. Map 106.
Voucher: Hardy 4869.
The small, blackish green plants with large,
conspicuous, hyaline perichaetia distinguish this
species from other Southern African taxa. The species
is also unique within Grimmiaceae because of the
stark contrast between the small, bistratose, muticose
vegetative leaves and the large, hyaline, piliferous
perichaetial leaves. In addition, the immersed,
cupulate capsules with rudimentary peristomes and
persistent annulus, which inflexes and thus narrows the
capsule mouth when dry, will help to identify the
species.
2. GRIMMIA
Grimmia Hedw., Spec. Muse. 75 (1801); Broth, in Natiirl. PflFam. 10: 306 (1924); Sim, Bryo.
S. Afr. 206 (1926). Lectotype species: G. plagiopodia Hedw., vide B.S.G., Bryol. Eur. 3: 99
(1845).
Plants small to medium, in dense tufts or cushions; saxicolous or terricolous. Stems irregu-
arly branched ; central strand present. Leaves frequently with hyaline point or awn. Laminal
cdls small above, at mid-leaf short-rectangular, sinuate; basal cells elongate, mostly straight-
walled.
Seta straight or curved; capsule immersed or exserted, smooth or 8-ribbed; peristome
teeth generally cleft above.
A widespread genus of 242 species, Grimmia is found throughout temperate and subtropical regions. The
genus is found in Southern Africa in association with rock substrates.
1 Capsules immersed ; leaves generally muticous or apex hyaline 1 . G. apocarpa
1 Capsules exserted; leaves with hyaline awn, short or long (sometimes absent):
2 Leaf lamina unistratose, margins thickened; capsule ribbed, seta frequently curved 2. G. pulvinata
2 Leaf lamina bistratose:
3 Leaves lanceolate to ovate-acuminate; basal leaf cells short-rectangular 3. G. ovalis
3 Leaves oval to ovate-lanceolate; lower juxtacostal cells frequently linear, basal cells quadrate
4. G. laevigata
1. Grimmia apocarpa Hedw., Spec.
Muse. 76 (1801); Broth, in Natiirl. Pflfam.
10:311 (1924); Sim, Bryo. S. Afr. 206 (1926).
Type: Germany, Dillenius s.n.
Grimmia depilis C. Mull., Syn. Muse. 1 : 778 (1849);
Broth, in Natiirl. PflFam. 10: 311 (1924). Type: Cape,
Philipstown, Pappe s.n.
Grimmia caffra C. Mull, in Hedwigia 38: 118
(1899); Broth, in Natiirl. PflFam. 10: 311 (1924).
Type: Orange Free State, Witteberg, Kadziberg,
Rehmann 1 30 (PRE!).
Grimmia boschbergiana C. Mull, in Hedwigia 38:
119 (1899); Broth, in Natiirl. PflFam. 10: 311 (1924).
Type: Cape, Somerset East, Boschberg, MacOwan
1877 (GRA!).
Grimmia oranica C. Miill. in Hedwigia 38: 119
(1899); Broth, in Natiirl. PflFam. 10: 311 (1924).
Syntypes: Orange Free State, Bethlehem, Rehmann
131 (PRE!); Kadziberg, Rehmann 132 (PRE!).
Plants medium-sized, forming tufts, dark
green, blackish below, occasionally reddish
or brownish, saxicolous. Stems erect to
inclined, 10-30 mm long, branched; in
section central strand small, inner cortical
cells in 4-5 rows, lax, outer cortical cells in
single row, stereids or substereids, yellowish.
Leaves spreading wet, little altered to erect
dry; ovate-acuminate, 1,5-2, 5 mm long,
frequently bistratose above; apex acute to
broadly acute, sometimes hyaline, infre-
quently with long, hyaline, toothed awn;
margins plane, entire, frequently bistratose,
occasionally multistratose above. Costa per-
current or extending into awn; in section
cells large, undifferentiated, incrassate. Upper
laminal cells quadrate to short-rectangular,
incrassate, frequently sinuate; basal cells
Grimmiaceae
275
short-rectangular to rectangular, occasionally
quadrate, walls smooth to sinuolate.
Autoicous. Perichaetia terminal, leaves
slightly larger, sheathing capsule. Seta short,
0,5 mm long, erect; capsule immersed, urn
ovoid, to 1,5 mm long, smooth; peristome
teeth triangular, perforated above, reddish
yellow; operculum rostrate; spores round,
12-15 pm, essentially smooth. Fig 81 : 19-29.
Almost cosmopolitan; in Southern Africa G.
apocarpa is frequently collected on rock, in grass-
lands of the Drakensberg of Lesotho, eastern Orange
Free State and Natal. The species is also collected in
the eastern Transvaal, Natal and the eastern, central
and southwestern Cape. Map 105.
Vouchers: Cholnoky 98; Crosby & Crosby 7944;
Hilliard & Bum 10612; Killick 4204; Mag ill 3369,
4680; Schelpe 7548; Schmitz 6971a.
This widespread species is variable and many of
its forms are present in Southern Africa. Plants col-
lected in the Karoo are generally small and little
branched with narrow, awned leaves, while plants
growing in stream-beds of the Drakensberg are
larger with broad leaves and generally lack awns or
hyaline apices. The species is easily distinguished
from other members of the family through its im-
mersed capsules, that are almost always present.
2. Grimmia pulvinata ( Hedw .) J. E. Sm.,
Engl. Bot. 24: 1728 (1807); Broth, in Natiirl.
PflFam. 10: 310 (1924); Sim, Bryo. S. Afr.
209 (1926). Type: Europe.
Fissidens pulvinatus Hedw., Spec. Muse. 158 (1801).
Fissidens pulvinatus var. africanus Hedw., Spec.
Muse. 159 (1801). Grimmia pulvinata var. africana
(Hedw.) Hook. f. & Wils. in Hook, f., FI. Nov. Zel.
2: 75 (1854). Type: Cape, Swartz s.n.
Grimmia leptotricha C. Mull, in Hedwigia 38: 120
(1899); Broth, in Natiirl. PflFam. 10: 310 (1924).
Type: Cape, Somerset East, Boschberg, MacOwan
s.n. (BM!).
Grimmia drakensbergensis Sim, Bryo. S. Afr. 209
(1926). Type: Natal, Symons sub Sim 9962 (PRE!).
Plants small, forming dense cushions,
blackish to yellow-green; saxicolous or
terricolous. Stems erect, 5-15 mm high,
branched below; in section central strand
small, inner cortical cells in 3-5 rows, large,
yellowish, outer cortical cells in 1-2 rows,
substereids, yellowish. Leaves appressed,
weakly spiralled dry, erect-spreading wet;
lanceolate to ovate-acuminate, 3, 0-3, 5 mm
long; apex acute to obtuse; margins plane,
entire, bistratose to multistratose above base.
Costa excurrent as a generally smooth,
hyaline awn, to 1 mm long, rarely percurrent ;
in section guide cells 2, large, ventral cells
absent, dorsal stereid band 3-4 cells thick,
dorsal surface cells large, incrassate. Upper
laminal cells short-rectangular, sinuate ; basal
cells long-rectangular to linear, walls smooth
or rarely sinuolate.
Autoicous. Perichaetia terminal; leaves
undifferentiated. Seta frequently curved, 2-3
mm long, yellowish; capsule exserted, in-
clined to pendulous, urn ovoid, 1 mm long,
8-ribbed dry; peristome teeth triangular,
2-3-cleft above, perforated below, weakly
papillose, red-yellow; operculum short-ros-
trate; spores round, 12-13 pm, smooth.
Fig. 82: 1-13.
Grimmia pulvinata is distributed worldwide. In
the Flora area it is most frequently collected in
grasslands of the Drakensberg of Natal and Lesotho
and in the Western Cape. Specimens have also been
collected in southern South West Africa/Namibia,
Natal, Orange Free State and the eastern, southern
and central Cape. Map 107.
Vouchers: Brenan M2738; Cholnoky 969;
Dieterlen 815; Edwards 659; Magill All 5, 4651;
Schelpe 7682; Schmitz 7870; Stokoe PRE-CH12774;
Van der Westhuizen & Deetlefs 26.
The leaf shape of G. pulvinata is variable and
sterile forms occasionally resemble G. apocarpa on
one extreme and G. ovalis on the other. An upper mid-
leaf section, however will reveal marginal thickenings
and only rarely, isolated areas of bistratose lamina.
The strongly ribbed, inclining or pendulous capsule
is also quite distinctive. In addition, the plants are
generally a lighter colour than other South African
species of Grimmia and have a greasy or oily appear-
ance when dry, making the leaves appear translucent.
Var. africana is occasionally separated (cf. Smith,
1978) on the basis of ovoid capsules and short-conical
opercula.
276
Grimmiaceae
Grimmiaceae
277
3. Grimmia ovalis ( Hedw .) Lindb. in Act.
Soc. Sci. fenn. 10: 75 (1871); Gangulee, Moss.
E. India, 1: 797 (1972); Grout, Moss FI. N.
Amer. 2: 33 (1933). Type: Europe, Saxony,
Hedwig s.n., 1792 (G).
Dicranum ovale Hedw., Spec. Muse. 140 (1801).
Grimmia commutata Hueb., Muse. Germ. 185
(1833), nom. illeg. ; Sim, Bryo. S. Afr. 210 (1926);
vide Sayre in Bryologist 54: 183 (1951).
Plants small to medium, olive-green to
blackish green, in dense cushions; saxicolous.
Stems erect, 5-20 mm tall, branching; in sec-
tion round, central strand small, inner
cortical cells in 3-5 rows, large, outer cortical
cells smaller, incrassate, yellowish. Leaves
appressed dry, spreading wet; ovate-acumi-
nate, 2-3 mm long; lamina bistratose above;
apex acute, generally with short, smooth,
hyaline awn; margins entire, frequently one
side weakly recurved. Costa percurrent to
short-excurrent; in section guide cells 2,
exposed ventrally, dorsal stereid or sub-
stereid band 1-2 cells thick, dorsal surface
cells large, incrassate. Upper laminal cells
quadrate, walls smooth ; basal cells rectangu-
lar, walls sinuolate.
Dioicous. Perichaetia terminal, leaves
oblong-acuminate, to 4 mm long. Seta
straight, 2,0-2, 5 mm long; capsule exserted,
urn ovoid to short-oblong, 1 ,0-1 ,5 mm long,
smooth; peristome teeth perforated or 2-3-
cleft above, weakly papillose, reddish yellow ;
operculum short-rostrate, beak ± oblique;
spores round, 9-12 pm, essentially smooth.
Fig. 82: 14-19.
A very widespread species; in Southern Africa
G. ovalis is concentrated in grasslands of the Drakens-
berg of Lesotho, Orange Free State and Natal. A few
specimens have also been collected in southern Natal
and the eastern, southern and western Cape. Map 106.
Vouchers: Edwards PRE-CH5437; Killick 4218;
Magill 4688; Schelpe 2116.
Where they overlap in the Drakensberg, this
species and G. pulvinata are superficially very similar.
Fertile specimens of G. ovalis are easily distinguished
by the smooth, erect capsules. Sterile specimens are
generally distinguished by their narrower leaves and
bistratose upper lamina.
4. Grimmia laevigata ( Brid .) Brid., Bryol.
Univ. 1: 183 (1826); Gangulee, Moss. E.
India 1 : 800 (1972); Scott & Stone, Moss. S.
Aust. 99 (1976). Type: Europe.
Campylopus laevigatus Burch., Mant. Muse. 76
(1819).
Grimmia campestris Burch, ex Hook., Musci Exot.
2: 129 (1819); Sim, Bryo. S. Afr. 208 (1926). Type:
Cape, Roggeveld, Burchell s.n.
Grimmia senilis Sim, Bryo. S. Afr. 207 (1926). Type:
Cape, Sim 9956 (PRE!).
Plants medium-sized, in loose tufts, dark
grey-green, hoary, or rarely yellow-green;
saxicolous or terricolous. Stems erect, to
20 mm tall, branching below; in section
central strand small, inner cortical cells in
4-5 rows, large, outer cortical cells in 2
rows, smaller, incrassate. Leaves appressed
dry, spreading wet; concave, lamina bistratose
above; ovate to broadly ovate, 3-4 mm long;
apex broadly acute, frequently hyaline,
extended into a long, spinose, hyaline awn, to
2 mm long, hyaline apical cells frequently
decurrent down margins of upper leaf ;
margins plane, entire or serrate at apex. Costa
broad below, quickly narrowing, weak above
mid-leaf but extending to apex; in distal
section guide cells 2, exposed ventrally,
dorsal stereid or substereid band small, 2-3
cells thick, dorsal surface cells larger, incras-
sate. Upper laminal cells quadrate to trans-
versely short-rectangular, somewhat longer at
basal angles; lower juxtacostal cells elongate,
merging with costa, rarely undifferentiated.
Dioicous. Perichaetia terminal; leaves
sheathing, oblong-aristate, to 4 mm long.
Seta short, straight, 1,0- 1,5 mm long,
yellowish; capsule emergent, urn elliptical,
1,0- 1,5 mm long, smooth, light brown;
peristome teeth cleft above, smooth, reddish
yellow; operculum short-rostrate; spores
round, 12-14 pm, smooth. Fig. 82: 20-26.
Fig. 82. — Grimmia pulvinata (1—13) : 1. habit, wet, x 1 ; 2. habit, dry, x 1 ; 3. habit, dry, x 5; 4. habit, wet,
x5; 5. leaves, x 10; 6. leaf in cross section, x435; 7. basal leaf cells at right margin, x 150; 8. cells at upper leaf
base (right side), x 150; 9. upper laminal cells at left margin, x 150; 10. upper laminal cells at right margin
(strongly incrassate form), xl50; 11. leaf apex, xl50; 12. part of capsule mouth with peristome teeth and
spores, x 270; 13. capsule and calyptra, x 5. G. ovalis (14-19) : 14. habit, wet, x 1 ; 15. habit, dry, x 1 ; 16. habit,
x5; 17. leaves, x25; 18. leaf in cross section, x220; 19. upper laminal cells, x290. G. laevigata (20-26): 20.
habit, wet, x 1 ; 21. habit, dry, x 1 ; 22. habit, x 5; 23-24. leaves, x 25; 25. leaf in cross section, x 220; 26. cells
at leaf base (left side), x 130. (1-13, van Zanten et al. 76091054; 14-19, Killick 4207; 20-26, Magill 4692).
278
Grimmiaceae
Grimmia laevigata has a worldwide distribution.
In Southern Africa, the species is frequently found on
rock or shallow soil in drier areas of the Cape,
Orange Free State, Lesotho and Natal. Map 108.
Vouchers: Esterhuysen 18785; Hilliard & Burtt
10664; Hardy 4313; Jacot Guillarmod 6122a; Lavranos
17001; Magill 4692; Relief PRE-CH12698; Schelpe
7815.
The leaf shape, broad costal base, and spinose
awn distinguish G. laevigata from other members of
the genus.
Insufficiently Known Species
Grimmia assurgens Shaw in Cape Monthly Mag.
17: 380 (1878). Type: Cape, Katberg, Shaw s.n.,
1869. The type has probably been destroyed. The
species is described as having “leaves papillose”
which lead Sim (1926) to suggest that it is related to
Tortula.
3. RACOMITRIUM
Racomitrium Brid., Mant. Muse. 78 (1819); Broth, in Natiirl. PflFam. 10: 312 (1924); Sim,
Bryo. S. Afr. 210 (1926). Lectotype species: R. canescens (Hedw.) Brid., vide Pfeiffer, Nom. 2:
907 (1874).
Plants medium to large, dark green to yellow-green, in large, loose cushions ; saxicolous.
Stems erect, with numerous short branches; central strand absent. Leaves lanceolate, fre-
quently with hyaline tip or awn. Laminal cells rectangular, sinuate or nodose.
Capsule exserted, erect, ovoid-cylindric, smooth; peristome 2-3-cleft above; operculum
rostrate; calyptra mitriform.
A very widespread genus of 79 species scattered on every continent, including Antarctica. In Southern
Africa Racomitrium is uncommon on rock in the southern and southwestern Cape Province and in isolated
localities in the Drakensberg of Natal and Lesotho.
1 Leaves with a long, hyaline awn; apical hyaline margin decurrent down leaf, toothed and papillose;
plants yellowish 1. R. lanuginosum
1 Leaves with short, hyaline awn or awn absent; leaf margins entire; plants dark green:
2 Alar cells differentiated, enlarged, quadrate, smooth-walled ; lower marginal cell walls sinuate ....
2. R. nigroviride
2 Alar cells not differentiated; lower marginal cells smooth-walled, occasionally in several rows. . . .
3. R. crispulum
1 . Racomitrium lanugiiiosum {Hedw.)
Brid., Mant. Muse. 79 (1819); Scott & Stone
Moss. S. Austr. 106 (1976). Type: Europe.
Trichostomum lanuginosum Hedw., Spec. Muse.
109 (1801).
Racomitrium hypnoides Lindb. in Ofvers. K.
VetenskAkad. Forh. 23: 552 (1866), nom. illeg.;
Sim, Bryo. S. Afr. 212 (1926).
Racomitrium lanuginosum var. pruinosum Wils. in
Hook, f., FI. Nov. Zel. 2: 76 (1854). Racomitrium
rpuinosum (Wils.) C. Miill. in Verh. zool.-bot. Ges.
Wien 19: 224 (1869). Type: New Zealand, North
Island, Colenso s.n.
Racomitrium incanum C. Miill. in Verh. zool.-bot.
Ges. Wien 19: 224 (1869). Syntypes: Cape, Table
Mountain, Ecklon s.n.; Thunberg s.n.
Plants large, in loose mats, yellow to
yellow-green, hoary; saxicolous. Stems erect
to inclined, 40-80 mm long, with numerous
short branches throughout; in section round,
central strand absent, inner cortical cells in
6 rows, medium-sized, smaller toward margin,
outer cortical cells in 1-2 rows stereids,
reddish. Leaves appressed with contorted
acumens dry, spreading wet; lanceolate to
Grimmiaceae
279
oblong-lanceolate, 3, 0-3, 5 mm long; apex
acuminate, hyaline, toothed and papillose,
hyaline region decurrent down margins;
base undifferentiated; margins plane to
recurved, toothed above, crenulate below.
Costa strong, ending in apex; in section
reniform, guide cells 6, large, strongly
thickened, exposed ventrally, dorsal cells
smaller, substereids. Laminal cells rectangu-
lar, strongly incrassate, nodose-sinuate; basal
cells larger, rectangular; basal marginal
cells smooth-walled, in single row, to 20
cells high.
Sporophyte not known from Southern
Africa. Fig. 83: 10-18.
This widespread species is generally locally
abundant in coastal mountain regions. In Southern
Africa, R. lanuginosum is found on rock or shallow
soils in the fynbos biome of the southern and south-
western Cape, especially on Table Mountain. Map
109.
Vouchers: Esterhuysen 15773; Pillans 4087; Sint
9287.
The large, hoary, yellow patches formed by this
moss are unmistakable. In addition the long-serrate,
hyaline hair-points are unique and denote the
Southern Hemisphere variety R. lanuginosum var.
pruinosum Hook. f. & Wils.
2. Racomitrium nigroviride (C. Mull.)
Par., Ind. Bryol. Suppl. 294 (1900); Broth, in
Natiirl. PflFam. 10: 313 (1924); Sim, Bryo. S.
Afr. 211 (1926). Type: Cape, Table Moun-
tain, Rehmann 139 (PRE!).
Grimmia nigroviridis C. Mull, in Hedwigia 38: 121
(1899).
Grimmia nigroviride var. robusticulum C. Mull, in
Hedwigia 38: 121 (1899). Racomitrium nigroviride var.
robusticulum (C. Mull.) Par., Ind. Bryol. Suppl. 294
(1900). Type: Cape, Devil’s Peak, Rehmann 139c
(NH!).
Grimmia pseudoacicularis C. Miill. in Hedwigia 38 :
120 (1899). Racomitrium pseudoacicularis (C. Miill.)
Par., Ind. Bryol. Suppl. 157 (1900). Type: Cape,
Drege s.n. (H-BR!).
Racomitrium defoliatum Dix. in Smithson, misc.
Colins 69(2): 16 (1918), vide De Sloover in Bull.
Jard. nat. bot. Belg. 47: 167 (1977). Type: Kenya,
western slope Mt Kenya, Mearns 1593 (BM; US).
Plants medium-sized, in tufts, blackish
to olive-green; saxicolous. Stems erect, 10-40
mm high; in section round, central strand
absent, inner cortical cells small, in 8-10
rows, thin-walled, becoming smaller toward
margin, thickened, yellowish, outer cortical
cells in 2-4 rows, small, stereids to sub-
stereids, red-yellow. Leaves erect dry,
recurved wet, weakly secund, lamina bistra-
tose becoming multistratose toward margins;
ovate-acuminate, 3, 5-4, 5 mm long; apex
acute, without hair-point, occasionally with
short hyaline tip; base auriculate; margins
incurved below, entire, without basal border
of smooth-walled cells. Costa strong, broad,
ending in apex; in section reniform, cells in
4-5 rows, large, incrassate, undifferentiated.
Laminal cells short-rectangular, incrassate,
nodose; alar cells forming distinct auriculate
group, enlarged, thin-walled, smooth-walled,
occasionally red-brown.
?Dioicous. Perigonia not seen. Peri-
chaetia terminal; leaves not differentiated.
Seta to 4 mm long, dark red; capsule ellip-
tical, 2 mm long; peristome teeth 2-3-cleft
to below middle, vertically striate below,
ornately papillose above, yellowish orange;
operculum rostrate, to 1 mm long; spores
round, 12-13 /mi, granulate. Fig. 83: 1-9.
Known from afro-alpine regions of eastern Africa
and the fynbos biome of Southern Africa, R. nigro-
viride is infrequently collected on the mountains of
the southwestern and southern Cape. Map 109.
Map 109. — • Racomitrium lanuginosum
x Racomitrium nigroviride
Vouchers: Crosby & Crosby 8179; Esterhuysen
18573, 21358; Primos PRE-CH9697; Thorne PRE-
CH3454.
The distinct alar group and absence of smooth-
walled border cells on the lower leaf margins separate
this species from R. crispulum. The unique leaf cells of
of Racomitrium distinguish R. nigroviride from other
related mosses.
280
Grimmiaceae
23
Grimmiaceae
281
3. Racomitrium crispulum {Hook. f. &
Wils.) Hook. f. & Wils. in Hook, f., FI. Nov.
Zel. 2: 75 (1854); Broth, in Natiirl. PflFam.
10: 313 (1924); Sim, Bryo. S. Afr. 212 (1926).
Type: Campbell Island, Menzies s.n. (BM).
Dryptodon crispulus Hook. f. & Wils. in Hooker,
Lond. J. Bot. 3: 544 (1844).
Racomitrium capense Lor., Moosstud. 163 (1864);
Broth, in Natiirl. PflFam. 10: 313 (1924). Type:
Cape, Ecklon s.n.
Grimmia austropatens C. Mull, in Hedwigia 38:
121 (1899), hom. illeg., non C. Mull, in Neum.,
Deutschl. Exp. Int. Polarforsch. 2: 316 (1890).
Racomitrium austropatens Broth, in Natiirl. PflFam.
10: 313 (1924). Type: Cape, Table Mountain, Reh-
mann 137 (NH!;PRE1).
Racomitrium drakensbergense Sim, Bryo. S. Afr.
212 (1926). Type: Natal, Symons sub Sim 8697
(PRE!).
Plants large, in dense tufts, olive-green
to blackish green; saxicolous. Stems erect,
20-70 mm tall; in section round, central
strand absent, inner cortical cells in 5-6
rows, incrassate, outer cortical cells in 1-2
rows, stereids, reddish. Leaves appressed dry,
recurved wet; ovate-acuminate, 3-4 mm long;
apex acute, rarely with short hyaline hair-
point; margins weakly recurved, entire; in
section lamina unistratose or bistratose
region restricted to marginal £ of lamina.
Costa strong, ending in acumen; in section
reniform, guide cells 6-8, large, incrassate,
exposed ventrally, dorsal cells smaller, incras-
sate. Laminal cells short-rectangular, strongly
incrassate, nodose; alar cells not forming
distinct group, reddish yellow; basal marginal
cells smooth-walled, to 25 cells high, 1 (-2)
cells wide.
Dioicous. Perichaetia terminal; leaves
not differentiated. Seta 3-4 mm long, yellow-
red; capsule cylindrical, 1,5 mm long; peri-
stome teeth irregularly 2-3-cleft to lower £,
papillose below, ornately papillose above,
yellow-orange; operculum rostrate, 1 mm
long; spores round, 17-18 um, granulate. Fig.
83: 19-27.
Widespread in the Southern Hemisphere, R.
crispulum is infrequently collected in the western,
southern and eastern Cape and a few localities in
Natal. Map 110.
Vouchers: Brenan M2771; Cholnoky 1011;
Esterhuysen 15774, 21326, 27049; Rourke 1688.
The differentiated basal marginal cells, bistratose
margins and generally unistratose lamina distinguish
Southern African specimens of R. crispulum.
Fig. 83. — Racomitrium nigroviride (1-9): 1. habit, wet, x 1 ; 2. habit, dry, xl;3. habit, x5;4. leaves, xlO;
5. leaf in cross section, x 125; 6. cells at leaf base (left side), x 170; 7. upper laminal cells, x435; 8. leaf apex,
X170; 9. part of capsule mouth with peristome teeth, x40. R. lanuginosum (10-18): 10. habit, wet, xl; 11.
habit, dry, xl; 12. habit, x5; 13. leaves, xlO; 14. leaf in proximal section, xl25; 15. leaf in distal section,
X 125; 16. basal cells at right margin, x435; 17. leaf apex, X40; 18. enlargement of awn (right margin), x 640.
R. crispulum (19-27): 19. habit, wet, x 1 ; 20. habit, dry, x 1 ; 21. leaves, x 10; 22. leaf in cross section, x 125;
23. cells at lower basal leaf margin, xl70; 24. upper laminal cells, x435; 25. leaf apex, xl70; 26. calyptra,
X5; 27. capsule, x5. (1-9, Thorne PRE-CH3454; 10-18, Boucher 3665; 19-27, Thorne PRE-CH6423).
283
INDEX TO FASCICLE 1
Acaulon C. Mull. 198, 202, 204, 206, 240
capense C. Mull 201
chrysacanthum Stone 199
leucochaete Stone 199, 201
muticum ( Hedw .) C. Mull 198, 201
recurvatum Magill 199
rufochaete Magill 201
sphaericum Shaw 201
Aloina (C. Miill.) Kindb 193, 206, 240
aloides ( Schultz ) Kindb 193
bifrons (De Not.) Delgadillo 193, 194
rigida sensu Sim 193
Andreaea Hedw 33, 35, 36
bistratosa Magill 37
nitida Hook. f. & Wils 35
' petrophila Fuemr 35
rupestris Hedw 33, 35, 36, 37
subulata Harv. ex Hook 33, 35
ANDREAEACEAE 33
Anisothecium Mitt.
natalense P. Varde 121
symonsii (Dix.) Broth 122
Anoectangium Schwaegr 179
assimilis Broth. & Wag 181
compactum Schwaegr 179
wilmsianum (C. Miill.) Par 181, 186
Anomobryum Schimp 120
Aongstroemia B.S.G 118
section Dicranella C. Miill 120
abruptifolia C. Miill 121
borgeniana Hampe 121
filiformis {P. Beauv.) Wijk & Marg 118, 120
gymnomitrioides Dix 110
julacea {Hook.) Mitt 120
longipes (Somm.) B.S.G 118
minuta Hampe 124
subsubulata Hampe ex C. Mull 121
transvaaliensis C. Miill 118
vulcanica (Brid.) C. Miill 118
ARCHIDIACEAE 71
Archidiella Irmsch.
dinteri Irmsch 72
Archidium Brid 71, 75, 81
subgenus Archidiella {Irmsch.) Snider 72
subgenus Archidium 72
acanthophyllum Snider 75
africanum Mitt 74
amplexicaule C. Miill 80, 81
andersonianum Snider 81
campylopodium C. Miill 78
capense Hornsch 78, 81
chrysosporum Schimp. ex Jaeg 80
compactum fo. tenerior C. Miill 78
dinteri {Irmsch.) Snider 72
ecklonii Hampe ex C. Miill 86
julicaule C. Miill 80
laterale Bruch ex Krauss 81
leptophyllum P. Varde 74
microthecium Dix. & P. Varde 74, 75
muelleranum Snider 77, 81
ohioense Schimp. ex C. Miill. 71, 74, 75
pellucidum Dix. ex Sim 74
phascoides Brid 71
rehmannii Mitt 77, 78, 81
subulatum C. Miill 77
Astomum Hampe 251
breutelianum Hampe ex C. Miill 84
nervosum (Hook.) C. Mull 86
pappeanum C. Miill 84
tetragonum (Hook.) Broth 250
Barbula Hedw. 213, 233, 236, 237, 239, 240, 246
section Asteriscium C. Miill 230
acutata C. Miill 213, 239
afrocaespitosa C. Miill 256
afrofontana (C. Miill.) Broth 246
afroinermis C. Miill 213
afroruralis C. Miill 219
anoectangiacea C. Miill 246
australasiae (Hook. & Grev.) Brid 232
brachyaichme C. Miill 215
brevimucronata C. Miill 216
caespitosa Schwaegr 256
calycina Schwaegr 240, 241, 243
chrysoblasta C. Miill 213
circinalis C. Miill 210
crinita Schultz 237, 239
deserta C. Miill 210
dimorpha C. Miill 233
dregeana C. Mull 193
erubescens C. Miill 215
erythroneura C. Mull 219
eubryum C. Miill 245, 246
eutrichostoma C. Miill 256
exesa C. Miill 215
flexicaulis C. Miill. 246
flexuosa (Hook.) Schultz 240
284
hildebrandtii C. Mull 215
hooker i Steud 241
homschuchiana Schultz 240
humilis Hedw 256
indica {Hook.) Spreng 243, 245
jamesonii Tayl 248
leptotortella C. Mull . 259
leucostega C. Mull 219
longipedunculata C. Mull 210
macowaniana C. Mull 215
microcalycina Magill 241, 243
muelleri B.S.G 219
natalensicaespitosa C. Mull 256
natalensis C. Mull 243
oranica C. Mull 215
orientalis (Web.) Broth 243
pagorum Milde 217
parvula (Hook. & Grev.) Spreng 209
perlinearis C. Miill 246
pertorquata C. Miill 246
pilifera (Hook.) Brid 237
porphyreoneura C. Miill. ex Vent 213
recurvata (Hook.) Schultz 209
rehmannii C. Miill 245
reticularia C. Mull 218
ruralis Hedw 219
schmidii C. Miill 215
torquatifolia Geh 213
torquescens Schimp. ex C. Miill 237
trichostomacea C. Miill 232
trivialis C. Mull 232
unguiculata Hedw 236
xanthocarpa C. Mull 235, 246
Bartramia Hedw 122
Blindia B.S.G 103
acuta {Hedw.) B.S.G 103
magellanica Schimp. ex C. Miill. 103, 104
Bruchia Schwaegr 106, 108, 111
brevipes Harv. ex Hook 108
eckloniana C. Miill 110
elegans (Homsch.) C. Miill 108
foveolata Magill 108, 110
rehmannii C. Miill 84
vogesiaca Schwaegr 106
BRUCHIACEAE 106
Bryobartramia Sainsb 269
novae-valesiae {Broth.) Stone & Scott . . 269, 270
robbinsii Sainsb 269
BR Y OBARTRAMIACEAE 269
Bryoerythrophyllum Chen 246, 248
afrorubellum (Broth. & Wag.) De Sloover . . . 248
jamesonii {Tayl.) Crum 248, 249
recurvirostrum {Hedw.) Chen 264, 248, 249
CALYMPERACEAE 156, 161
subfamily Octoblepharoideae 161
Calymperes 163, 164
levyanum Besch 166
lonchophophyllum Schwaegr 163
nashii Williams 164
rabenhorstii Hampe & C. Miill 164
tenerum C. Miill. var. edamense Fleisch. 164, 166
victoriae Dix 166
Campylopus Brid. 138, 139, 143, 154
section Microcampylopus C. Miill 137
altovirescens (C. Mull.) Par 146
ampliretis (C. Miill.) Par 140, 142
angustinervis Dix 138
atroluteus (C. Miill.) Par. .. . 147, 148, 150, 154
aureoviridis (C. Miill.) Schimp. ex Par 154
bartramiaceus (C. Miill.) Ther. ex Broth. 148, 150
basalticolus (C. Miill.) Par 146
bequaertii Ther. & Nav 143, 144, 147
bewsii Sim 152
bicolor (C. Mull.) Wils 150
var. ericeticola (C. Miill.) Dix 150
bryhnii Broth 146
capensis Broth 142
catarractilis (C. Mull.) Par 148
chlorotrichus (C. Miill.) Par 142
delagoae (C. Miill.) Par 140, 142
echinatus Sim 150
edwardsii Sim 140
flexuosus {Hedw.) Brid. 138
griseolus (C. Miill.) Par 152
heteroneurum Ther 140
imerangae orth. var 146
inandae (C. Miill.) Par 142
inchangae (C. Miill.) Par 143, 146, 147, 150
introflexus {Hedw.) Brid. 138, 150, 151, 152, 153
laevigatus Brid 277
lepidophyllus (C. Miill.) Jaeg 150
leptotrichaceus (C. Miill.) Par 148
leucobasis (C. Mull.) Par 150
longescens (C. Miill.) Par 146
nanotenax (C. Miill.) Par 146
nanus C. Miill 121
olivaceonigricans (C. Miill.) Par 140
pallidus Hook.f. & Wils 147, 150
perfalcatus Broth 142
perpusillus Mitt 138
pilifer Brid. 152, 153
285
polytrichoides De Not 152
procerus (C. Mull.) Par 153, 154
pseudobicolor C. Mull, ex Ren. & Card. 150
pseudojulaceus Sim 144
pulvinatus (C. Miill.) Par 147
purpurascens Lor 152
purpureoaureus (C. Mull.) Par 152
serridorsus (C. Mull.) Par 146
simii Schelpe 144
stenopelma (C. Miill.) Par 142, 143, 147
stramineus {Mitt.) Jaeg 143
subchlorophyllosus C. Miill. ex Rabenh. 144, 146
symonsii Sim 152, 153
tenax (C. Miill.) Broth 142
trichodes Lor 152
var. perlamellosus Dix 152
weisiopsis (C. Miill.) Par 150
Ceratodon Brid. 94, 246
purpureus ( Hedw .) Brid. 94, 96
stenocarpus Bruch & Schimp. ex C. Miill. . . 94
Chorisodontium {Mitt.) Broth 154
aciphyllum {Hook, f & Wils.) Broth 155
dicranellatum {Dus.) Roiv 155
falcatum Magill 154
Cladophascum Dix 110, 111
gymnomitrioides {Dix.) Dix. ex Sim 110
Conomitrium Mont 39
capensis C. Miill 68
erosulum C. Miill 58
gracile Hampe ex C. Mull 51
julianum (Cand.) Mont 67
pseudo serratum C. Miill 61
scleromitrius Besch 55
Crossidium Jur 195, 197, 198, 206, 240
apiculatum Magill 195, 197
spiralifolium Magill 197
squamiferum ( Viv.) Jur 195
Cynodontium Schimp.
capillaceum Hedw 98
Desmatodon Brid. 209, 210
convolutus {Brid) Grout 209, 210, 212
latifolius {Hedw.) Brid. 209
longipedunculatus (C. Miill.) Magill 210, 212
recurvatus (Hook.) Mitt 209
Diaphanophyllum Lindb.
vallis-gratiae Lindb 93
DICRANACEAE 69, 89, 105, 156, 161
subfamily Dicranelloideae 118
subfamily Dicranoideae 126
subfamily Leucobryoideae 156, 161
subfamily Trematodontoideae 106, 111
Dicranella {C. Miill.) Schimp. .. 69,70,89,120,
121, 122, 126
abruptifolia (C. Miill.) Par 121
borgeniana (Hampe) Jaeg 121
minuta (Hampe) Jaeg 124
nana Wint 121
rigida Dix. ex Sim 122, 124
subsubulata (C. Miill.) Jaeg 121
symonsii Dix 121, 122
Dicranodontium B.S.G.
chlorotrichus (C. Mull.) Par 142
laxitextum Broth. & Bryhn 146
per falcatum (C. Miill.) Par 142
Dicranoloma (Ren.) Ren 130, 133
billardieri (Brid.) Par 130, 132, 133
entabeniense Magill 132
nitidulum (C. Miill.) Par 133
Dicranum Hedw.
section Chorisodontium Mitt 154
altovirescens C. Mull 146
ampliretis C. Miill 140, 148
atroluteus C. Miill 148
aureoviridis C. Miill 154
bartramiaceus C. Mull 148
basalticolus C. Miill 146
billardieri Brid 132
catarractilis C. Mull 148
chlorotrichus C. Miill 142
chrysobasilare C. Mull 133
delagoae C. Miill 140
filiforme P. Beauv 118
griseolus C. Miill 152
inandae C. Miill 142
inerangae C. Miill 146
intro flexum Hedw 150
lepidophyllus C. Miill 150
leptotrichaceus C. Miill 148
leucobasis C. Miill 150
longescens C. Miill 146
nano tenax C. Mull 146
nanum (C. Miill.) C. Miill 121
nitidulum C. Miill 133
olivaceonigricans C. Miill 140
ovale Hedw 277
perfalcatum C. Miill 142
procerus C. Miill 153
pulvinatus C. Miill 147
purpureoaureus C. Miill 152
purpureus Hedw 94
286
rehmannii C. Mull 136
serridorsus C. Mull. 146
sprengelianum C. Mull 134
stenopelma C. Miill 142
tenax C. Miill 142
weisiopsis C. Miill 150
zeyheri C. Miill 134
Didymodon Hedw 232, 233
afer (C. Mull.) Broth 233
afrorubellus Broth. & Wag. ex Dix 248
capensis Spreng 210
ceratodonteus (C. Miill.) Dix 233, 235
dimorphum (C. Miill.) Broth 233
fragilis Hook. & Wils 258
knysnae Rehm. ex Sim 235
lingulatum (Sim) Magill 184
perrevolutus P. Varde 235
pottsii Dix 233
rigidulus Hedw 233
squarrosus Hook. 189
xanthocarpus (C. Miill.) Magill 235, 236, 246
Distichium B.S.G 98
capillaceum ( Hedw .) B.S.G 98
DITRICHACEAE 83, 89
Ditrichum Hampe 89, 93
amoenum ( Thwait . & Mitt.) Par 89, 93
brachypodum (C. Miill.) Broth 92, 93
brevifolium (C. Miill.) Broth 92
capense (C. Miill.) O. Kuntze 93
cylindricarpum (C. Miill.) F. Miill 90
difficile (Dub.) Fleisch 93
flexifolium Hampe 93
glaucescens (Hedw.) Hampe 96
homomallum (Hedw.) Hampe 89
hymenodontium Dix 89, 92, 93
punctulatum Mitt 90, 92
spirale Dix 89, 92
strictum (Hook. f. & Wils.) Hampe 90
vallis-gratiae (Lindb.) Hampe 93
Dryptodon Brid.
crispulus Hook. f. & Wils 281
Eccremidium Hook. f. & Wils 88
exiguum (Hook. f. & Wils.) Wils 88, 89
Encalypta Hedw 173
ciliata Hedw 173, 174, 176
vulgaris Hedw 174, 176
ENCALYPTACEAE 173, 269
Ephemerum Hampe 201
Eucladium B.S.G 186
verticillatum (Brid.) B.S.G 186
Fissidens Hedw 39, 63, 69, 70, 108
subgenus Aneuron Kindb 41, 44
subgenus Fissidens 43, 44
subgenus Octodiceras (Brid.) Broth 67
subgenus Pachyfissidens (C. Miill.) Kindb. ... 67
section Aloma C. Miill 62
section Areofissidens C. Miill. 43, 44, 58
section Crenularia C. Miill. 61, 62
section Crispidum C. Miill. 62, 63
section Fissidens 46, 47
section Pachyfissidens C. Miill 67
section Semilimbidium C. Miill 54, 58
section Serridium C. Miill. 63
aciphyllus Dix 53, 54
amblyophyllus C. Miill 65
androgynus Bruch ex Krauss 46
aristatus Sim 50
arnellii P. Varde 44
asplenioides Hedw 63, 65
bifrons Schimp. ex C. Miill 45, 57, 58
bocarangensis P. Varde 46
borgenii Hampe 58, 59, 61
var. obtusifolius Dix 59
breutelii Schimp. ex C. Miill 48, 50
brevisetus Sim 55
bryoides Hedw 39, 46, 47
calochlorus Dix 55
capensis (C. Mull.) Broth 68
capriviensis Magill 45
curvatus Hornsch 51, 53, 68
cuspidatus C. Miill 51
cymatophyllus C. Miill 62
enervis Sim 41
erosulus (C. Miill.) Par 58, 59
eschowensis Broth. & Bryhn 54
fasciculatus Hornsch 65, 67
flaccidus Mitt 46
fontanus (B. Pyl.) Steud. 67, 68
glaucescens Hornsch 62, 63, 65, 67
gracilis (Hampe) Jaeg 51
gueinzii C. Miill 46
haakonii Broth. & Bryhn 58
hoeegii P. Varde 47
hyalobasis Dix. ex Sim 55
ischyrobryoides C. Miill 50
julianus (Cand.) Schimp 67
lanceolatus Hampe 62
laxifolius Hornsch 51
linearicaulis Broth. & Bryhn 58
longulus C. Miill. 68
macleanus Shaw 68
macowanianus C. Miill 48
malacobryoides C. Miill 46
marginatus Schimp. ex C. Miill 50
287
megalotis Schimp. ex C. Miill 47
microandrogynus Dix 55, 57
minutiretis Dix 48
mucronatus Schimp. ex C. Mull 62
nitens Rehm. ex Salm 62
opacifolius Mitt 54
papillifolius Dix 54
parvilimbatus Sim 57
pauperrimus C. Mull 68
pectinidens Dix 57
perpaucifolius Dix. ex Sim 57
plumosus Hornsch 63, 65, 67
procerior Broth. & Bryhn 62
pulvinatus Hedw 275
var. africanus Hedw 275
pseudoserratus (C. Miill.) Jaeg 61, 62
pycnophyllus C. Miill 51, 68
pygmaeus Hornsch 45, 57, 58, 68
rehmannii C. Miill 62
remotifolius C. Miill 46
rufescens Hornsch 47, 48, 50
rutenbergii Par 68
scleromitrius ( Besch .) Broth 55
simii Schelpe 50
splachnifolius Hornsch 44, 45, 58
stellenboschianus Dix. ex Sim 51, 53
submarginatus Bruch ex Krauss 54
subobtusatus C. Miill 58, 59, 61
subremotifolius C. Miill 46
urceolatus Wag. & Dix 54
usambaricus Broth 43
vittatus Hook. f. & Wils 48
wageri Dix. ex Wag 44
zuluensis Broth, eft Bryhn 62
FISSIDENTACEAE 39
Fontinalis Hedw.
antipyretica Hedw 68
juliana (Savi) Savi ex Cand 67
Goniomitrium Hook. & Wils 270
Grimmia Hedw 271, 274, 275
apocarpa Hedw 35, 246, 274, 275
assurgens Shaw 278
atrovirens J. E. Sm 209
austropatens C. Miill 281
boschbergiana C. Miill 274
caffra C. Miill 274
campestris Burch, ex Hook 277
commutata Hueb 277
depilis C. Miill 274
drakensbergensis Sim 275
laevigatus ( Brid .) Brid. 277, 278
leptotricha C. Mull 275
nigroviridis C. Miill 279
var. robusticulum C. Miill 279
oranica C. Miill 274
ovalis ( Hedw .) Lindb 275, 277
plagiopodia Hedw 274
pseudoacicularis C. Miill 279
pulvinata (Hedw.) J. E. Sm 275, 277
var. africana (Hedw.) Hook. f. & Wils. . . . 275
senilis Sim 277
GRIMMIACEAE 271, 274
Gymnostomum Nees & Hornsch. ... 182, 183, 184
aeruginosum J. E. Sm 183, 184
bewsii Dix 183, 184, 186
calcareum Nees & Hornsch 183
dimorphum (C. Miill.) Sim 233
gracile (Wag. & Dix.) Dix 233
gracile (R. Br.) Hook 235
involutum Hook 228
julaceum Hook 120
lingulatum Rehm. ex Sim 183, 184
pottsii (Dix.) Sim 233
recurvirostre Hedw 182
wageri Schelpe 235
Gyroweisia Schimp.
amplexicaulis Sim 221
latifolia Dix 221
Holomitrium Brid. 129
affine Card. <ft Ther 129
cylindraceum (P. Beauv.) Wijk & Marg. var.
cucullatum (Besch) Wijk & Marg. 129, 130
perichaetiale (Hook.) Brid. 129
vaginatum (Hook.) Brid. var. cucullatum
(Besch.) C. Mull 129
Husnotiella Card. 221
latifolia (Dix.) Zander & Magill . . . 221, 222, 224
obtusifolia (Broth.) Zander 222
plicata Magill 222
revoluta Card. 221
Hydrogonium (C. Miill.) Jaeg 236, 237
afrofontanum (C. Miill.) Hilp 246
ehrenbergii (Lor.) Jaeg 236
Hymenostomum R. Br 251, 262
brachycarpum (C. Miill.) Par 263
cucullatum (C. Miill.) Kindb 266
eurybasis Dix 267
humicola (C. Miill.) Par 266
latiusculum (C. Mull.) Par 267
microstomum (Hedw.) R. Br 262
opacum Wag. & Dix 243
288
Hymenostylium Brid.
ceratodonteum (C. Mull.) Broth.
crassinervium Broth. & Dix
recurvirostrum ( Hedw .) Dix 182,
xanthocarpum {Hook.) Brid.
Hyophila Brid 225,
afrophaea (C. Mull.) Wamst
atrovirens (C. Mull.) Broth
baginsensis C. Miill
basutensis Sira
cyathiforme (Dix.) Sim
involuta {Hook.) Jaeg 228,
javanicum (Nees & Blum.) Brid.
plicata Mitt
zeyheri (Hampe) Jaeg
Hypodontium C. Miill
dregei {Hornsch.) C. Miill 170,
pomiforme {Hook.) C. Miill.
var. macowanianum C. Miill
Leptodontium (C. Miill.) Hampe ex Lindb. 187,
brachyphyllum Broth. & Ther 191,
longicaule Mitt 191,
var. microruncinatum {Dus.) Zander
squarrosum (Hook.) Hampe ex Lindb
viticulosoides {P. Beauv.) Wijk & Marg
Leptotrichum Hampe ex C. Miill.
brachypodum C. Miill
brevifolium C. Miill
capense C. Miill
LEUCOBRYACEAE 156,
Leucobryum Hampe
acutifolium {Mitt.) Card 156,
glaucum {Hedw.) Schimp.
gracilifolium Dix
gueinzii C. Miill
isleanum Besch
madagassum Besch 158,
parvulum Card.
perfalcatum Sim
rehmannii C. Miill
Leucoloma Brid.
albocinctum Ren. & Card.
bifidum {Brid.) Brid.
chrysobasilare (C. Miill.) Jaeg 133,
haakonii Broth. & Bryhn
rehmannii (C. Miill.) Par 134, 136,
sprengelianum (C. Miill.) Jaeg 134, 136,
syrrhopodontioides Broth
zeyheri (C. Miill.) Kindb 134, 136, 137
zuluense Broth. & Bryhn 133
Leucoperichaetium Magill 271
eremophilum Magill 271, 273, 274
LEUCOPHANACEAE 161
Lophiodon Hook. f. & Wils.
strictus Hook. f. & Wils 90
Microcampylopus (C. Miill.) Fleisch 137, 138
nanus (C. Miill.) Broth 121, 138
perpusillus {Mitt.) Broth 138
pusillus C. Miill 121
subnanus (C. Miill.) Fleisch 137
Microdus Schimp. ex Besch 124
minutus {Hampe) Besch 124
Moenkemeyera C. Miill 39
mirabilis C. Miill 39
scleromitria (Besch.) P. Varde 55
NANOBRYACEAE 69
Nanobryum Dix 69, 70
dummeri Dix 69, 70
gladiolum {Mitt.) Bizot 70
Neckera Hedw.
viticulosoides P. Beauv 189
Octoblepharum Hedw 156,161
albidum Hedw 161, 163
Octodiceras Brid 39, 67
capensis (C. Miill.) Jaeg 68
fissidentoides Brid 39
fontanum (B. Pyl.) Lindb 67
julianum (Cand.) Brid 67
Oreoweisia {B.S.G.) De Not 127
bruntonii {Sm.) Mild. 127
erosa (C. Miill.) Kindb 127, 129
serrulata (Funck) De Not 127, 129
torquescens {Brid.) Wijk & Marg 129
Oxystegus {Limpr.) Hilp 259
cylindricus {Brid.) Hilp 259, 260
Phasconica C. Miill. 251, 253
balansae C. Miill. 251
lorentzii C. Miill 251
tisserantii P. Varde 251, 253
Phascum Hedw 198, 202, 204, 206
cuspidatum Hedw 202
elegans Hornsch 108
exiguum Hook. f. & Wils 88
leptophyllum C. Miill 202, 204
182
233
183
183
182
228
260
228
230
235
170
230
228
224
260
170
172
172
172
189
192
192
191
198
189
92
92
93
161
156
158
156
158
156
158
159
158
158
158
133
137
133
134
136
137
137
136
289
nervosum Hook 86
peraristatum C. Mull. 202, 204
splachnoides Homsch 208
tetragonum Hook 250
Pleuridium Rabenh 83, 86, 108
breutelianum (Hampe) Jaeg 84
ecklonii (C. Mull.) Snider 86
nervosum ( Hook .) Mitt 86, 88
papillosum Magill 88, 89
pappeanum (C. Miill.) Jaeg 84, 86, 88
subulatum ( Hedw .) Rabenh 83
POLYTRICHACEAE 198
Pottia ( Reichenb .) Fuernr 204, 206, 209, 210
afrophaea C. Miill 260
ceratodontea C. Miill 233
drummondii ( Wils.) Willis 209
macowaniana C. Miill 207, 208
maritima (R. Br. ter.) Broth 209
namaquensis Magill 206, 207
splachnoides ( Homsch .) Broth 207, 208, 209
subplanomarginata Dix 207, 208
truncata (Hedw.) B.S.G 204
verrucosa Rehm. ex Warnst 207, 208
zeyheri Hampe ex C. Miill 260
POTTIACEAE 127, 177, 236, 269
subfamily Pottioideae 178
subfamily Trichostomoideae 249
tribe Barbuleae 227
tribe Leptodontieae 186
tribe Pleuroweisieae 179
tribe Pottieae 192
Pseudocrossidiu M Williams 213, 239, 240
Pterygoneurum Jur 197, 240
cavifolium Jur 197
mdcleanum Warnst 197, 198
Racomitrium Brid. 278, 279
austropatens Broth 281
canescens (Hedw.) Brid. 278
capense Lor 281
crispulum (Hook.f. & Wils.) Hook. f. & Wils. 279, 281
defoliatum Dix 279
drakensbergense Sim 281
hypnoides Lindb 278
lanuginosum (Hedw.) Brid. 278, 279
var. pruinosum Wils 278, 279
incanum C. Miill 278
nigroviride (C. Miill.) Par 279
var. robusticulum (C. Miill.) Par 279
pruinosum (Wils.) C. Miill 278
pseudoacicularis (C. Miill.) Par 279
Saelania Lindb 96
caesia (P. Beauv.) Lindb 96
glaucescens (Hedw.) Broth 96
Schistidium Brid 246
Schistomitrium Dozy & Molk.
acutifolium Mitt 156
SELIGERIACEAE 103
Semibarbula Herz. ex Hilp 236, 237
indica (Hook.) Herz. ex Hilp 236, 243
orientalis (Web.) Wijk & Marg 243
Skitophyllum B. Pyl 39
fontanum B. Pyl 67
Sphaerangium Schimp.
capense (C. Mull.) Jaeg 201
SPHAGNACEAE 23
Sphagnum L 23
africanum Welw. & Dub 28, 29
austromolle C. Miill 31
balfourianum Warnst 27
beyrichianum Warnst 32
capense Homsch 31, 32
var. austromolle (C. Miill.) Warnst 31
var. mollissimum (C. Miill.) Warnst 31
var. multiporosum Warnst 32
convolutum Warnst 29
coronatum C. Miill 29
var. cuspidatum Warnst 29
fimbriatum Wils. in Hook 24, 26
fluctuans C. Mull 29
laceratum sensu Garside 26
marlothii Warnst 27
marginatum Warnst 29
var. convolutum Warnst 29
var. diversifolium Warnst 29
mollissimum C. Miill 31
obtusiusculum Warnst 26
oligodon C. Miill 28
var. bachmannii Warnst 28
var. beyrichii Warnst 28
oxycladum Warnst 29
palustre L 23
panduraefolium C. Mull 29
pappeanum C. Miill 26
perichaetiale Hampe 27
pycnocladulum C. Miill. 29, 31, 32
var. fuscescens Warnst 29
var. viride Warnst 29
rehmannii Warnst 28
strictum Suit, subsp. pappeanum (C. Miill.)
Eddy 26, 27
290
transvaaliense Wamst
truncatum Hornsch
Sporledera Hampe
rehmannii (C. Mull.) Kindb
Syrrhopodon Schwaegr
dregei Homsch
erectifolius C. Miill
gardneri {Hook.) Schwaegr
gomesii P. Varde
obliquirostris C. Miill.
perichaetialis Bruch ex Krauss
pomiformis (Hook.) Hampe ex C. Mull.
uncinifolius C. Miill. 167,
Tetrapterum Hampe ex Jaeg 250,
australe Hampe
tetragonum {Hook.) Andrews 250,
Thysanomitrion Schwaegr
delagoae (C. Miill.) Broth
inandae (C. Mull.) Broth
richardii (Brid.) Schwaegr
transvaaliense Herz. & Dix. ex Sim
Timmiella {De Not.) Limpr 253,
anomala {B.S.G.) Limpr
cameruniae Broth
pelindaba Magill
Tortella {Lindb.) Limpr 255,
afrocaespitosa (C. Mull.) Broth
caespitosa (Schwaegr.) Limpr 255,
calycina (Schwaegr.) Dix
eutrichostomum (C. Miill.) Broth
fragilis {Hook. & Wils.) Limpr 255, 258,
humilis {Hedw.) Jenn 255, 256,
natalensicaespitosa (C. Miill.) Broth
petrieana Sim
rufiseta (C. Miill.) Broth 255,
tortuloides (Sull. & Lesq.) Broth 255,
xanthocarpa (C. Miill.) Broth 255, 256,
Tortula Hedw. 212, 213, 215, 218, 239, 240, 246,
afroruralis (C. Miill.) Broth
ammonsiana Crum & Anderson 216,
atrovirens (J.E. Sm.) Lindb.
australasiae Hook. & Grev. ...
bifrons De Not
brachyaichme (C. Mull.) Broth.
brevimucronata (C. Miill.) Broth.
brevitubulosa Broth
deserta (C. Miill.) Broth
erubescens (C. Miill.) Broth. . .
erythroneura (C. Miill.) Broth.
eubryum (C. Miill.) Broth
exesa (C. Miill.) Broth 215
flexuosa Hook 240
hildebrandtii (C. Miill.) Broth 215
indica Hook 243
irregularis Sim 216
leucostega (C. Miill.) Broth 219
longipedunculata (C. Miill.) Broth 210
macowaniana (C. Miill.) Broth 215
muelleri Hook. f. & Wils 219
muralis Hedw 213, 215
oranica (C. Miill.) Broth 215
pagorum {Mild.) De Not 216, 217, 218
papillosa Wils. ex Spruce 218
parvula Hook. & Grev 209
pilifera Hook 237
var. longifolia Sim 237
porphyreoneura (C. Miill.) Townsend ... 213, 239
princeps De Not 218, 219
recurvata Hook 209
reticularia (C. Miill.) Broth 218
ruralis {Hedw.) Gaertn., Meyer & Scherb. 219, 221
schmidii (C. Miill.) Broth 215, 216
subtorquati folia Dix 210
subulata Hedw 212
torquatifolia (Geh.) Dix 213
trachyneura Dix 219, 221
Trachycarpidium Broth.
novae-valesiae Broth, ex Roth 269
Trematodon Michx Ill, 117
subgenus Gymnotrematodon C. Miill. ex
Broth Ill
subgenus Trematodon Ill
aequicollis Ren. & Card 115
africanus Wag. & Dix 115, 117
divaricatus Bruch ex Krauss 114,115
flexifolius C. Miill 115
intermedius Welw. & Duby 112, 114
ligulatus Rehm. ex Roth 117
longicollis Michx Ill, 115, 117
mayottensis Besch 117
pallidens C. Miill 115
paradoxus Hornsch 114, 115
pillansii Dix. ex Sim 112
Trichostomopsis Card. 230, 232, 233
australasiae {Hook. & Grev.) Robinson . . 232, 233
crispifolia Card. 230
trivialis (C. Miill.) Robinson 232
Trichostomum Bruch 251, 260
section Hydrogonium C. Miill 236
afrofontanum C. Miill 246
afrum C. Miill 233
28
29
84
166
170
167
166
167
169
170
172
169
251
250
251
138
140
142
138
140
254
253
254
253
259
256
256
240
256
259
258
256
256
256
256
262
278
219
218
209
232
193
215
216
216
210
215
219
245
291
atrovirens Rehm. ex C. Mull
brachydontium Bruch ex F. A. Muell. 253, 260,
convolutum Brid
cyathiforme Dix
cylindricum (Brid.) C. Mull
difficile Dub
glaucescens Hedw
lanuginosum Hedw
leiodontium C. Miill
orientalis Web
rehmannii Sim 255,
rufisetum C. Mull
squarrosum (Hook.) Schwaegr
tortuloides Sull. & Lesq 255,
xanthocarpum Schimp. ex C. Mull
Triquetrella C. Miill.
papillata {Hook. f. & Wils.) Broth
strictissima Rehm. ex C. Miill
tristicha (C. Mull.) C. Mull 187,
Tristichiopsis C. Miill.
mirabilis C. Miill
Tristichium C. Miill
lorentzii C. Miill.
mirabile (C. Miill.) Herz
Weisiopsis Broth 224, 225, 227,
anomala {Broth. & Par.) Broth
involuta Magill 225,
plicata {Mitt.) Broth 224, 225
pulchriretis Dix 225
Weissia Hedw 251, 253, 262, 263, 267, 268
subgenus Hymenostomum {R. Br.) Kindb. . . 262
subgenus Weissia 262
subgenus Oreoweisia B.S.G 127
brachycarpa C. Miill 263
controversa Hedw 262, 263, 265, 266
var. pillansii Schelpe 263
cucullata C. Miill. 266, 267
cylindricus Bruch ex Brid 259
dieterlenii Ther 263, 265, 266
erosa Hampe ex C. Miill 127
gracilis Wag. & Dix 233
humicola C. Miill 263, 265, 266
latiuscula C. Miill 267, 268
linguaelata Shaw 265
oranica C. Miill 265
pomiformis Hook 172
recurvirostris Hedw 248
vallis-gratiae C. Miill 265
viridula Hedw. ex Brid 265
var. brachycarpa (C. Miill.) Dix 263, 266
var. longifolia Broth. & Wag 265
vulcanica Brid 118
Zygodon Hook. & Tayl.
simii Dix 189
tristichus C. Mull 187
wilmsianum C. Miill 181
228
262
209
170
259
93
96
278
262
243
256
255
189
256
255
187
188
187
188
99
99
99
99
230
224
227
Vol. 12: Portulacaceae, Basellaceae, Caryophyllaceae, Illecebraceae, Cabombaceae, Nymphaeaceae, Cerato-
phyllaceae, Ranunculaceae, Menispermaceae, Annonaceae, Trimeniaceae, Lauraceae, Hernandia-
ceae, Papaveraceae, Fumariaceae
Vol. 13: Brassicaceae, Capparaceae, Resedaceae , Moringaceae, Droseraceae, Roridulaceae , Podostemaceae,
Hydrostachyaceae (Published 1970). Price: R10,00. Overseas: R12,00. Post free
Vol. 14: Crassulaceae
Vol. 15: Vahliaceae, Montiniaceae, Escalloniaceae, Pittosporaceae, Cunoniaceae, Myrothamnaceae, Brunia-
ceae, Hamamelidaceae, Rosaceae, Connaraceae
Vol. 16: Fabaceae: Part 1: Mimosoideae (Published 1975). Price: R13,50. Overseas: R16,75. Post free
Part 2: Caesalpinioideae (Published 1977). Price: R16,00. Overseas: R20,00. Post free
Papilionoideae
Vol. 17: Geraniaceae, Oxalidaceae
Vol. 18: Linaceae, Erythroxylaceae, Zygophyllaceae, Balanitaceae, Rutaceae, Simaroubaceae, Burseraceae,
Ptaeroxylaceae, Meliaceae, Aitoniaceae, Malpighiaceae
Vol. 19: Polygalaceae, Dichapetalaceae, Euphorbiaceae, Callitrichaceae, Buxaceae, Anacardiaceae, Aquifo-
liaceae
Vol. 20: Celastraceae, Icacinaceae, Sapindaceae, Melianthaceae, Greyiaceae, Balsaminaceae, Rhamnaceae,
Vitaceae
Vol. 21 : Tiliaceae, Malvaceae, Bombacaceae, Sterculiaceae
Vol. 22: Ochnaceae, Clusiaceae, Elatinaceae, Frankeniaceae, Tamaricaceae, Canellaceae, Violaceae, Flacour-
tiaceae, Turneraceae, Passifloraceae, Achariaceae, Loasaceae, Begoniaceae, Cactaceae. (Published
1976.) Price: R8,60. Overseas: RIO, 75. Post free
Vol. 23: Geissolomaceae, Penaeaceae, Oliniaceae, Thymelaeaceae, Lythraceae, Lecythidaceae
Vol. 24: Rhizophoraceae, Combretaceae, Myrtaceae, Melastomataceae, Onagraceae, Trapaceae, Haloraga-
ceae, Gunneraceae, Araliaceae, Apiaceae, Cornaceae
Vol. 25: Ericaceae
Vol. 26: Myrsinaceae, Primulaceae, Plumbaginaceae, Sapotaceae, Ebenaceae, Oleaceae, Salvadoraceae, Loga-
niaceae, Gentianaceae , Apocynaceae. (Published 1963.) Price: R4,60. Overseas: R5,75. Post free
Vol. 27: Part 1: Periplocaceae, Asclepiadaceae (Microloma-Xysmalobium)
Part 2: Asclepiadaceae (Schizoglossum-Woodia)
Part 3 : Asclepiadaceae (Asclepias-Anisotoma)
Part 4: Asclepiadaceae ( Brachystelma-Riocreuxia ). (Published 1980.) Price: R4,50. Overseas: R6,00.
Post free
Asclepiadaceae (remaining genera)
Vol. 28 : Cuscutaceae, Convolvulaceae, Hydrophyllaceae, Boraginaceae, Stilbaceae, Verbenaceae, Lamiaceae,
Solanaceae, Retziaceae
Vol. 29: Scrophulariaceae
Vol. 30: Bignoniaceae, Pedaliaceae, Martyniaceae, Orobanchaceae, Gesneriaceae, Lentibulariaceae, Acantha-
ceae, Myoporaceae
Vol. 31: Plantaginaceae, Rubiaceae, Valerianaceae, Dipsacaceae, Cucurbitaceae
Vol. 32: Campanulaceae, Sphenocleaceae, Lobeliaceae, Goodeniaceae
Vol. 33: Asteraceae
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11