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Section B. Biological Sciences
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Saprotrophic fungi of the Guinea–Congo Region
T. Laessøe, L. Ryvarden, Roy Watling and A. J. S. Whalley
Proceedings of the Royal Society of Edinburgh. Section B. Biological Sciences / Volume 104 /
January 1996, pp 335 - 347
DOI: 10.1017/S0269727000006187, Published online: 05 December 2011
Link to this article: http://journals.cambridge.org/abstract_S0269727000006187
How to cite this article:
T. Laessøe, L. Ryvarden, Roy Watling and A. J. S. Whalley (1996). Saprotrophic fungi of
the Guinea–Congo Region. Proceedings of the Royal Society of Edinburgh. Section B.
Biological Sciences, 104, pp 335-347 doi:10.1017/S0269727000006187
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Proceedings of the Royal Society of Edinburgh, 104B, 335-347, 1996
Saprotrophic fungi of the Guinea-Congo Region
T. Laessee
Institute for Sporeplanter , University of Copenhagen, Denmark
L.
Ryvarden
Department of Botany, University of Oslo, Norway
Roy Watling
Royal Botanic Garden, Edinburgh EH3 5LR, U.K.
and
A. J. S. Whalley
School of Biomolecular Sciences, John Moore's University, Liverpool, U.K.
Synopsis
A window onto the vast array of saprotrophic fungi found in West Central Africa with particular
reference to the Cameroon attempted in this paper. After an overview (Watling) three diverse groups are
taken as examples of this array: xylariaceous fungi (Laess0e, Whalley & Watling); polypores (Ryvarden
& Watling) and lepiotoid fungi (Watling). Some comparisons are made with other tropical forests. This
approach contrasts a lignicolous group in the ascomycetes with one in the basidiomycetes and contrasts
terricolous lepiotoid agarics with polyporaceous terricolous and lignicolous fungi.
Overview
Saprotrophic fungi in West Central Africa, in common with those of other rain
forests of the world, are an integral part of the vegetation but their full diversity is
not fully understood. They undoubtedly play a most important role in the rapid
recycling of nutrients although this has yet to be measured quantitatively. Knowledge
of the fungi involved is rather poor and based on limited observations; either
collections at the turn of the century, when the European empires were expanding
and explorers and missionaries were sending material back to Europe, or those made
after the Second World War when the colonies so formed were being developed and
exploited. Little organised collecting was carried out but nevertheless some sort of
picture, however cloudy, is emerging. With the collecting being so patchy it is difficult
to judge whether some of the information available is meaningful.
Hennings (1895, 1897, 1901, 1905) described 62 species of Marasmius from West
Africa (all but four new to science) based on material collected by Zenker, Dusen
etc.,
but only four species of Mycena. Is this because of the difficulty of drying
Mycena in the rain forest compared to relative ease with which basidiomes of
Marasmius can be preserved, or does it reflect a real preponderance of the latter?
Some fungi are undoubtedly under-represented in present day collecting but in
general Marasmius, the xylariaceous fungi and the polypores are more obvious
336 T. Laessoe et al.
constituents than many of the other groups and some have very spectacular basidi-
omes,
e.g. the bright red Donkia sanguinea Maas Geest. and Favolaschia zenkeri-
ana Singer.
Beeli (1928) described ten species of Marasmius from the Congo, whilst Singer
(1965) recorded 66 from the same region. In Britain the ratio of Marasmius to
Mycena is 30:102 whereas in a single transect in Korup National Park, Cameroon
at least 60 taxa of Marasmius were recognised compared to fewer than 12 species
of Mycena.
Lentinus tuber-regium (Fr.) Fr. is widespread and common and L. velutinus Fr.,
L.
squarrulosus Mont, and L. brunneofloccosus Pegler are all found in Korup.
Hennings (1895) described 13 species of Panus and Lentinus, 8 of them new, although
these are now classified in several often quite unrelated genera. Pegler (1972)
recorded 15 species from the Congo, 5 of which appear in Hennings and all appear
in Bresadola (1890): L. crinitus, (L.:Fr.) Fr., L. anthocephalus (Lev.) Pegler (as
L.
braunii Bres.), L. exilis (Klotzsch) Bres., and L. tanghama Lev., synonymous with
L.
sajor-caju (Fr.) Fr. Lentinus spp. are an important source of food for tribal people
(Watling 1991) and local Africans report the eating of
lignicolous,
pleurotoid agarics
by primates, especially chimpanzees.
Members of the families Bolbitiaceae, Coprinaceae, Hygrophoraceae, and
Strophariaceae are infrequent within the Korup rain forest although they are
widespread in the disturbed areas outside the forest. Surprisingly the number of
the Pluteaceae are similar to that of warm temperature countries, likewise the
Lepiotaceae, which is a colourful and prominent member of the flora. The members
of the last family are a feature of one of the specialist 'parts' below. Romagnesi
(1957) has described 12 members of the Entolomataceae from Zaire although
Romgnesi & Gilles (1979 & 1980) record 179 from Gabon and the Ivory Coast;
several are now recognised in Cameroon.
If all these collections are taken into account a very substantial fungal flora results.
It has in part been illustrated and documented in Flore Iconographique des
Champignons du Congo published by the Jardin Botanique de l'Etat; Bruxelles, later
retitled the Flore Illustree des Champignons d'Afrique Centrale.
Certain trends amongst the saprophytic species are explored below. In addition
widespread taxa such as Lentinus tuber-regium are found from West and East Africa
to Australasia and the New World, whereas L. brunneoflocculosus seems to be
confined to the present region (Ghana to Zaire). Some of the aphylloporoid fungi
also show similar distributional patterns, viz. in Podoscypha, a genus of prominently
lignicolous fungi. P. bolleana (Mont.) Boid., common in West Africa, is probably
limited to tropical Africa whereas P. involuta (Klotzsch apud Fr.) Imazeki and
P.
mellisei (Berk. & Sacc.) Pat. are also found in SE Asia. The former is common
throughout the tropics except the New World.
Boidin (1966) includes amongst the ten species of Cymatoderma and Podoscypha
with which he deals all the Cameroon species including P. mellisei which is rare
outside Malaysia. Cymatoderma elegans Jungh and C. dendriticum (Pers.) Reid are
found in the Cameroon.
The resupinates found at Korup include Hydnophlebia chrysorhiza (Torr.) Parm.
and Phlebia chrysocreas (Berk. & Br.) Burdsall which are pan tropical species,
Hyphodontia gossypina (Parm.) Hjortst. on the other hand forms a species-pair with
H. brevidens (Pat.) Ryv. from S. America in a similar way to Cystidiodontia isabellina
Saprophytic fungi 337
(Bert. & Br.) Hjortst. & Ryv. and C. artocreas (Berk. & Curt.) Hjortst. & Ryv.
Resupinates are found on twigs, branches and larger woody debris on the forest
floor. However, some debris is often retained in the canopy and only revealed by
climbing amongst the trees, examining lianas pulled down in a tree-fall and searching
areas of tree-fall. Species which occur in such habitats include Hyphodontia nespori
(Bres.) Erikss. & Hjortst. and unidentified species of both Trechispora {Wat. 22779
in E) and Hyphodontia (Wat. 22792 in E); similar results have been found in the
forest canopy in Campo province, Cameroon (Nunez & Ryvarden, personal com-
munication). This woody resource is similar to that recorded for marasmioid fungi
where hammocks collecting leaves above head height are formed, a phenomenon
described from South America (Hedger 1989) but recognised also in the Cameroon
forest. The fusing together of woody debris on the forest floor in a parallel way is
also a feature of the rain forest. The taxa involved include Theleporus calcicolor
(Sacc. & Syd.) Ryv., Hydnophlebia chrysorhiza and a flame orange Steccherinum sp..
The Hymenochaetaceae, comprising corticioid forms as well as poroid resupinate
and pileate forms, are important members of the forest community, breaking down,
depending on the species involved, everything from twigs to standing, old, emergent
giants, e.g. P. alliardii (Bres.) Ryv. on Lecomtedoxa klaineana (Pierre ex Engl.)
Dubard and Phellinus durissimus (Lloyd) Ryv. on Oubanguia alata Bak. f. Some of
the trees show extensive decay with large cavities or hollow trunks. These act as
suitable resting places for nocturnal mammals whose dung is an important source
of nitrogen for the forest.
Termites are also an important component of the rain forest community, feeding
on leaf and woody material, and contributing to the nitrogen cycle. Associated with
the termites, is a rather specialised group of agarics placed in the genus Termitomyces.
Four species have been recorded from Korup in the present survey, viz.
T. microcarpus (Berk. & Br.) Heim, T. clypeatus Heim, T. globulus Heim & Goos.
and T. striatus (Beeli) Heim. In addition T. medius Heim has been found at Kumba
and T. letestuei (Pat.) Heim at Bipinde based on Lepiota discopus Henn. (as
L.
clavipes Henn., Zenker 1568 in E). Heim (1952) records 10 species from Congo.
The genus is known from other parts of Africa and SE Asia but not from Australia
nor South America (Heim, 1977).
'Gasteromycetes', although not common, are widespread and include examples of
all the major groups. For example Cyathus limbatus Tul., Sphaerobolus stellatus
Tode: Pers., Geastrum schweinitzii (Berk. & Curt.) Zeller, Lycoperdonpusillum Batsch
& L. polymorphii Vitt., and Scleroderma dictyosporum Pat. have all been recorded
from Korup along with some potentially ectomycorrhizal species (see Bucyk, Thoen
& Watling this volume). Dictyophora duplicata (Vent.) Desv. is frequently cited from
West Central Africa (see Demoulin & Dring 1975; Dissing & Lange 1962, 1963
& 1964).
Larger ascomycetes, other than members of the Xylariaceae considered below,
are rather uncommon. Cookeina sulcipes (Berk.) Kuntze on woody debris is probably
the most conspicuous, although collections of Phaedropezia epispartia (B. & Br.) Le
Gal and a species close to Galactinia auriformis (Pat.) Le Gal, both described from
Zaire (Le Gal 1960), have recently been found in Korup.
One of the most spectacular fungi of this part of Africa is the hyphomycete
Sarophorum palmicola (Henn.) Seifert & Samson (anamorph of Penicillopsis clavaria-
eformis Solms-Laubach) which grows on hard, fallen fruits of forest trees. It is very
338 T. Laessoe et al.
polymorphic and many growth forms agreeing with several of the names introduced
by Hauman (1936) have been seen in Korup, e.g.
Penicillopsis longissima
Hauman
and P. microsequoia Hauman: see Samson & Seifert (1985). P. africana Seifert &
Samson and its anamorph Stilbodendron was also found (Wat. 22167-70 & 72).
The accounts of specific groups which follow rely heavily on recent material
collected in Cameroon, especially during the period
1989-91,
by one of us (RW),
and during 1991 by L. Ryvarden. Additional material from elsewhere in the Guinea-
Congo lowland rain forest area has been incorporated. The vegetation of that part
of Korup National Park, SW Cameroon in which fungi were collected has been
described by Gartlan et al. (1986).
Xylariaceous fungi
The Xylariaceae are a prominent feature of the fungus flora in the lowland rain
forest of Korup. The fruiting bodies of many species are conspicuous through their
size,
and were found in both the ascigenous state and, perhaps more frequently in
the case of
Xylaria,
in their anamorphic state. Representative taxa occurred in many
different habitats ranging from twigs and branches to woody legume pods, matted
leathery leaves, and large fallen trunks. Xylaria exhibited the greatest range in
substrata. Twenty six species belonging to six different genera have been collected
from Korup. Many of these species have also been recorded by Dennis from the
Congo (Dennis 1962 & 1963) and Sierra Leone (Dennis 1958), but in Korup there
are notable absentees and fewer taxa overall. The Korup collections are almost
equally divided between the erect strap-like or club-shaped stromata typical of
Xylaria, and the hemispherical, pulvinate or effused forms found in Hypoxylon sensu
lato and its close allies.
The most widespread species appear to be Kretzchmaria clavus (Fr.) Sacc. (5
collections), Xylaria globosa (Spreng ex Fr.) Mont. 9 collections and as many
sightings and X.
ianthinovelutina
(Mont.) Fr.. Many of the collections of
X.
globosa
were colonised by Nectriopsis puiggarii (Speg.) Samuels which until now has not
been reported from Africa although recorded from South America, New Zealand,
the Philippines and Sri Lanka (Samuels 1988). One specimen of X. globosa was
colonised by an unidentified member of the genus
Ceraceomyces
(Wat. 22978 in E;
Hjortstam, personal communication). Ustulina deusta (Hoffm: Fr.) Lind occurred
on logs close to those colonised by K. clavus. We are unable to confirm whether
they are conspecific, as suggested by Ko et al. (1982) but we certainly believe them
to be congeneric. A monographic treatment of Kretzschmaria is needed.
Sarcoxylon cf. compunctum (Jungh.) Cooke was found on three occasions but, in
common with a number of collections of other taxa, was immature indicating that
mid-January to mid-April is not favourable for good teleomorph development in
these taxa. Over 58 collections of Xylaria were made during this period of which 27
(46%) were either completely immature or conidial. Even by May Sarcoxylon strom-
ata were only just beginning to ripen. Anamorphic stromata of Xylaria are much
easier to locate due to their copious production of white conidia. One collection of
Xylaria (Wat. 21229) is new to science and awaits a formal description. Another
taxon related to X. corniformis (Fr.) Fr. also appears to be new.
Poles from the forest are cut by local people for building shelters, for fashioning
staffs, and for the construction of handrails over water-courses. On these structures
Saprophytic
fungi 339
Biscogniauxia nummularia
(Bull, ex St. Amans) O. Kuntze (represented by three
varieties) and
H.
citriforme
Whalley, Hammelev
&
Taligoola were found. Hypoxylon
citriforme
has only recently been described from Nigeria (Whalley et al. 1988) and
is also known from Malaysia (Whalley & Jones, unpublished). A large spored form
occurs in Papua New Guinea; this taxon also belongs in
Biscogniauxia
(Van der
Gucht & Whalley 1992).
Camillea tinctor
(Berk.) Laessoe, J. D. Rogers & Whalley
was also collected on cut poles along with H. stygium (Lev.) Sacc; the latter was
also found in undisturbed forest. These observations although based on limited
material, are partially supported by data from elsewhere. Thus Laessoe et
al.
(1989)
found species of
Camillea
to be inhabitants of sun-exposed sites such as natural
clearings and man-made fellings. Taligoola
&
Whalley (1976) reported
Biscogniauxia
and
Hypoxylon
to be fungi of drier sites in three Ugandan forests and reported that
H.
{Biscogniauxia) nummularium
var. merrillii (Bres.) Mill, and H. thouarsianum
(Lev.) Lloyd are primary colonisers of fairly sound, clean material either on the
forest floor or on branches still attached to trees.
Hypoxylon
rubiginosum
(Pers.: Fr.) Fr. was represented in Korup by two distinct
varieties, the type variety and var.
dieckmannii
(Theiss.) Miller which is widespread
throughout the tropics (Miller 1961). Xylaria
globosa,
X.
obovata
(Berk.) Fr. and
Sarcoxylon cf. compunctum were all found on large, very rotten trunks; the
Sarcoxylon
often occurred in association with other members of the family. Laessee
et
al.
(1989) noted that
Xylaria
and
Kretzschmaria
often occurred in wet rather than
dry situations.
Xylaria ianthino-velutina
was commonly found on the pods of caesalpi-
noid legumes and is undoubtedly a common inhabitant of leguminous pods and
other woody fruits throughout the tropics (Rogers 1979; Laessee, unpublished). No
species of
Rosellinia
Ces. & De Not. were found; they are apparently rare in rain
forests (Rogers et
al.
1987).
It is perhaps surprising that the majority of species recorded occur throughout
the tropics but it is encouraging to discover that species occur in similar habitats in
other tropical regions. Thus it appears that their distribution is very much linked to
environment. As the available data are still very restricted, and far from adequate,
it is difficult to make any generalisations. However, a comparison of both genera,
and the number of species in some of these genera, with those reported from South
America
(e.g.
Dennis 1970) indicate that the Xylariaceae are much better represented
there than in either Africa or Asia (Whalley 1993).
Polypores
The polypores are a striking component of the lowland rain forest ranging from
small basidiomes scarcely
10
mm across to enormous basidiomes reaching over
300 mm in diameter. The full spectrum of fruit-body morphology is also found,
from centrally stipitate to dimidate/pileate and totally resupinate forms. Basidiomes
may be soft and fleshy
(Laetiporus sulphureus
(Fr.) Murr.), flexible and slightly tacky
(Gloeoporus thelephoroides
(Hook.) Gunn.), both of which show extensive attack by
invertebrates, tough and hard
{Phellinus
spp.), or leathery
Amauroderma
spp.). Even
the toughest basidiomes show distinctive burrows caused by insect activity. Both
annual and perennial life-strategies are found amongst the 67 species recorded from
Korup, SW, Cameroon and the 123 species from Campo province in SE Cameroon.
There appear to be many similarities between the two sites.
340 T.
Laessoe
et al.
In Korup the genus
Microporus
was extremely common, represented by six species
which grow on woody debris on the forest floor, ranging from twigs to fallen logs.
Equally common and widespread was
Nigropoms vinosus
(Berk.) Murr. with basidi-
omes rather more difficult to see in the gloom than the equally common, white,
Gloeoporus
thelephoroides.
These and members of the genus
Phellinus,
of which ten
are known from Korup alone, are the dominant members of the flora. The more
interesting finds, however, have been amongst the less well-distributed taxa.
Laetiporus
sulphureus
is interesting in that it is also a familiar member of the
W European polypore flora; L.
percicinus
(Berk. & Curt.) Gilbert, known also from
the New World, is a gigantic terrestrial species resembling in colour, shape and
possibly smell, a large
Bankera
until turned over. At Korup, where it is known from
two sites, it appears to fruit from a perennial mycelium.
Piptoporus soloniensis
(Fr.)
Pilat, a rare cosmopolitan species found in the Korup rain forest resembles
L.
sulphureus and was previously known as P.
pseudosulphureus
Long (see
Schumacher & Ryvarden 1981).
The other large polypores include several members of the
Ganoderma lucidum
group, G.
australe
(Fr.) Pilat and
Amauroderma
spp.. A.
kwiluensis
(Beeli) Ryv. is
particularly interesting. Until the Cameroon collections were
made,
and these number
over eight with as many sightings, it was only known by a single collection
from Zaire (Ryvarden & Johansen 1980). Other interesting taxa are -
Perenniporia
contraria
(Berk. & Curt.) Ryv. (very rare, known also from Cuba, Rwanda and
Kenya), P.
ochroleuca
(Berk.) Ryv. and Schizopora
flavopora
(Cke.) Ryv. (both
cosmopolitan).
Many of the other polypores found are known from other areas of the globe and
appear to be fairly widespread. Thus
Pycnoporus sanguineus
(Fr.) Murr. is known
from tropical Africa and from Central and South America; it is replaced by the
reportedly different,
P.
coccineus
(Fr.) Murr. in Australia and
P.
cinnabarinus
(Jacq.:
Fr.) Karst. in Europe.
Trametes
and Lenzites spp. in the West African rain forest
appear to be pantropical, e.g. T.
vespacea
(Pers.) Ryv. and T.
elegans
(Fr.) Fr.. The
closely related genus
Coriolopsis
also contains pantropical taxa, e.g. C. byrsina
(Mont.) Ryv. and
C.
caperata
(Berk.) Murr.
(=Datronia),
although
C.
badius
(Berk.)
Murr., which is common in Asia and Australia, is apparently rare in Africa.
Coltricia
spathulata (Hook.) Murr. and C.
cinnamomea
(Pers.) Murr. are known from all
areas of the globe, the latter even in north to central Europe. Their appearance
beneath ectomycorrhizal tree species raises the possibility that they might be ectomy-
corrhizal; certainly in Korup C.
spathulata
was associated with
Uapaca.
Aurificaria
indica
(Mass.) Reid is related to
Coltricia
and has been found in Korup. It is usually
recorded from Asia but there is a record from Kenya (Ryvarden & Johansen 1980).
In contrast
Phylloporia chrysita
(Berk.) Ryv. is found in
S
America and The southern
USA, in addition to Asia and West Africa.
Several
Phellinus
spp. collected recently in West Central Africa are either pantrop-
ical, e.g.
P.
senex (Nees & Mont.) Imaz. and
P.
sublamaensis
(Lloyd) Ryv. or known
from the Indo-Malaysian area, e.g. P. extensus (Lev.) Pat. P. gilvus (Schw.) Pat.
and
P.
ferreus (Pers.) Bourd. & Galz. are also pantropical and the latter even reaches
Europe. P. noxius (Corner) Cunn., known from Sierra Leone to Tanzania, is an
important parasite.
On basidiomes of a
Phellinus
sp. the fruiting structures of the corticioid
Phlebia
ryvardenii
Hallen. & Hjortst. were collected; it has previously only been collected in
Saprophytic
fungi 341
Spain, Sweden and Eastern Soviet Union. In Korup P.
allardii
(Bres.) Ryv. and
P.
durissimus
(Lloyd) Ryv. are the commonest taxa. In addition to
Gloeoporus
thelephoroides
(Hook.) Cunn., a pantropical species, the commonest species in the
lowland rain forest are
Theleporus calcicolor
(Sacc. & Syd.) Ryv. (Grammothelaceae:
African & Indo-Malaysian), Porogramme albocincta (Cke. & Mass.) Lowe
(Polyporaceae: pantropical, recorded Ethiopia to Malawi) and
Rigidoporus
microp-
orus (Fr.) Overeem. (Polyporaceae: tropical to subtropical parts of Asia, America
& Africa). R.
ulmarius
(Fr.) Imaz., very familiar to European mycologists, is known
from the tropics including W Africa, and R.
biokensis
(Lloyd) Ryv. is pantropical
but, although recorded from Korup, is rare in Africa. Porogramme albocincta
appeared to be confined to small diameter
Uapaca
stems.
Two pantropical species, Megasporoporia cavernulosa (Berk.) Ryv. and
Tinctoporellus epimiltinus
(Berk. & Br.) Ryv. are widespread in African lowland
rain forest. In contrast Loweoporus
roseoalbus
(Jungh.) Ryv. is palaeotropical.
Porodisculus pendulus
(Schw.) Murr., found only once in the Korup rain forest, is
delicate, attractive and almost
Panellus-like;
it is pantropical, although rather rare.
Because of its size it may be under-recorded. Three species of
Microporellus
have
been collected in Cameroon, M. obovatus (Jungh.) Ryv. a pantropical species,
M.
collybiaeformis
(Beeli) Ryv. only previously known from Zaire, and an as yet
unassigned, probably new, taxon.
It appears that the polypores of the Guinea-Congo Region include both pantrop-
ical and palaeotropical elements as well as species whose distribution extends to
Europe and N America. Only a few species are restricted to Africa and these are
distributed through to East Africa or widely south of the Sahara. There are some
hints of a neotropical element but more collecting is required. What little collecting
has been carried out supports the thesis that brown-rotting genera are characteristic
of the boreal zone. Many of the polypores in the rain forest surrounding the Bight
of Biafra are structurally complex including a suite of di- and tri-mitic taxa. This
parallels observations from the better known East African flora.
Lepiotoid fungi
In classical texts the genus Lepiota is characterised by the presence of a ring, either
membranous or friable, lack of a volva, white or pale-coloured spore-print, and free
gills.
With the use of microscopic characters it soon became apparent that this was
an artificial grouping and now several independent genera are recognised. They
include
Cystolepiota,
Leucoagaricus, Leucocoprinus,
Macrolepiota
and
Sericeomyces;
in addition there is a more carefully circumscribed residual genus Lepiota.
Representatives of all these genera have been found in the Guinea-Congo lowland
rain forest area. Chamaemyces and
Pseudobaeospora
are smaller genera, as is
Verrucispora,
which was first found in the Cameroon, based on material collected
by Zenker at Bipinde (No.
2171
in E). It is known from Central and East Africa
and was originally placed in
Inocybe,
(as Lflavofusca) by Hennings (1901) undoubt-
edly because of the shape of the basidiospores (Kuyper 1990). It was not found
during the present survey. The true mushrooms
(Agaricus
spp.), of which there are
several taxa in the lowland rain forest, are closely related to Lepiota although they
have a chocolate, umber or even red or purple tinged spore-mass (Heinemann 1956,
1957 & 1986).
342 T.
Laessoe
et al.
Lepiotoid fungi are terricolous and may be found in a variety of habitats from
open grassland to mature woodland. One is immediately struck in the tropics by
the large number and variety of lepiotoid basidiomes. Subsequently, it comes as a
surprise when the numbers of taxa amongst what might be a hundred or so basidi-
omes found in a single day in the rain forest proves to be quite small. Indeed the
number of taxa involved is less than the range found in Europe (101), and of the
same order as that of the British Isles (61), a country which is not considered
notable for its extensive collection of Lepiota sensu lato recorded at its forays
especially as one travels north.
In the rain forest a range of lepiotoid fungi are found in damp shade, margins of
tree falls and tracks, under or around fallen trunks, or along damp banks and
overhangs of streams. Some idea of the diversity in the area can be judged by the
accounts of Beeli (1926) and Heinemann (1967, 1970, 1973, 1977 & 1980) from
Zaire where 66 taxa are covered. The number of taxa recorded is similar to that in
East Africa (Pegler 1977; 48 with 13 possible additions), Lesser Antilles (Pegler
1983;
40), and Sri Lanka (Pegler 1986; 39). A number of species eg. Lepiota
subincarnata,
L.
epicharis,
Leucocoprinus
birnbaumii,
L.
caepestipes
and Sericeo-
myces serena
are common to all four areas (Table 1). These
figures
however, rely on
a few sporadic sightings over many years coupled with at least one lengthy study
as an adjunct to some other profession. This contrasts with Britain where inten-
sive collecting has been carried out over 150 years in an enormous variety of
geological and vegetational types.
Surprisingly Henning (1895 et subseq.) described only 9 species from Cameroon
but this is probably because many taxa to be found there are fragile and difficult to
preserve and transport to Europe. Judging from recent collections the numbers of
taxa will parallel those for Central Africa. Of the 37 taxa, (Table 1), based on 58
collections found during the present study in the Korup rain forest the greatest
numbers are assignable to
Leucocoprinus
and Lepiota Sect.
Ovisporae
(Table 2).
Hundreds of
Leucocoprinus
basidiomes appear throughout the forest after early
rains,
often carpeting the floor or growing on or under rotten algal covered trunks.
Two taxa were dominant during April 1990, viz. Leucocoprinus
fragillissimus
(Rav.
apud Berk. & Curt.) Pat. and L.
birnbaumii
(Corda) Singer (Lepiota lutea (Bolt.:
Seer.) Godfrin; =L. flos-sulphuris (Schnizlein) Cejp). L.
birnbaumii
is common in
the British Isles and Europe associated with greenhouse-grown plants. It was
recorded in the first instances from Botanic Gardens, e.g. Edinburgh, Kew, Paris,
Munich, Prague and is now of ten recorded accompanying pot plants in nurseries
and homes.
Lepiota subincarnata
J. Lange is known from both Europe and East Africa (Pegler
1977).
Several of the taxa found in Cameroon have been found in East Africa either
as newly described species, L.
rubiginosa
Pegler and
Leucoagaricus rhodocephalus
Pegler, or more widespread taxa e.g. L.
citrinophylla
(Berk. & Br.) Sacc. and
L.
pseudogranulosa
(Berk. & Br.) Sacc. Other species are common to both Zaire
and Cameroon, e.g. L.
violaceovelutina
Beeli, L.
testacea
Beeli, L.
purpureoimbricata
Beeli, L,
echinella
var.
congolensis
Beeli and the common L.
coactila
Beeli.
All the main genera and sections of lepiotoid fungi are found in the rain forest
except
Macrolepiota
and
Chlorophyllum
which are only known from worked land
adjacent to the forests. In addition several very interesting taxa have been collected
in Korup. These include a
Leucoagaricus
with a bronze-coloured spore-print and a
Saprophytic fungi
343
Table 1. Lepiotoid fungi collected in Korup National Park, S.W. Cameroon in the
period 1984-91 with notes on their distribution
Cystolepiota (=Lepiota Sect. Micacei J. Lange.)
Lepiota
Sect. Cristatae
L.
ceramogenes (Berk. & Br.) Sacc.
Known from E. Africa; described from Sri
Lanka.
L.
epicharis (Berk. & Br.) Sacc.
Known from E. Africa, and a variety from
Venezuela; described from Sri Lanka.
L.
micropholis (Berk. & Br.) Sacc.
Known from E. Africa; described from Sri
Lanka. Recorded as alien from Europe.
L.
montageni var.
Known from East & Central Africa*.
L.
rubiginosa Pegler
Described from East Africa.
Additional unidentified species - 1
Sect. Ovisporae
L.
coactila Beeli
Described from Central Africa.
L.
lepidosphora (Beck. & Br.) Sacc.
Described from Sri Lanka.
L.
cf. lilacea Bres.
Known from Europe & S. America.
L.
cf. phaeosticta Morgan
Described from the New World & East Africa.
L.
pwpureoimbricata Beeli
Described from Central Africa.
L.
subincarnata J. Lange
Described from Europe; known in East and
West Africa.
L.
testacea Beeli
Described from Central Africa*
L.
violaceovelutina
Described from Central Africa
Additional unidentified species - 4
Sect. Stenosporae
L.
citrinophylla (Berk. & Br.) Sacc.
Known from E. Africa: described from Sri
Lanka.
Additional unidentified species - 1
L.
echinella var. congolensis Beeli
Variety described from C. Africa; type variety
from Europe.
L.
pseudogranulosa (Berk. & Br.) Sacc.
Known from East Africa; described from Sri
Lanka.
L.
aff. sistrata (Fr.) Quel.
Species described from Europe.
Additional unidentified species - 1
Leucoagaricus
L.
rhodocephalus Pegler
Described from East Africa.
L.
fuligineus Pegler
Described from East Africa.
Sericeomyces (=Lepiota sect. Sericellae Ktthn.)
S. serena (Fr.) Heinem.
Described from Europe. Known from Central
& East Africa and Sri Lanka.
S. viscidulus Heinem.
Described from Central Africa.
Also see L. eriphaea (Berk. & Br.) Sacc.
assignable here.
Known from East Africa; described from Sri
Lanka.
Leucocoprinus
L.
birnbaumii (Corda) Singer
Australasia, India, Africa & New World also
recorded as an alien in Europe.
L.
brebissonii (God. ) Locq.
Known in disturbed habitats in Europe & New
World.
L.
cepaestipes (Sow.: Fr.) Pat.
Known in disturbed environments in Europe &
New World.
L.
flava (Beeli) Heinem.
Described from Central Africa. + closely related
taxon.
L.
fragillissimus (Rav.) Pat.
Known from East and Central Africa, & Sri
Lanka; described from southern USA.
Leucocoprinus sp.: olivaceous pileus & stipe;
yellow gills.
*See Heinemann (1936).
344 T.
Laessoe
et al.
Table 2. Number of taxa of lepiotoid fungi in tropical agaric floras
Lepiota
Sect. Ovisporae
Sect. Echinatae
Sect. Stenosporae
Sect. Cristatae
Sect. Lepiota
Chlorophyllum
Cystolepiota
Leucoagaricus
Leucocoprinus
Macrolepiota
Sericeomyces
TOTAL
Korup:
Cameroon
10(12)
1(1)
2(3)
5(6)
2(2)
1(1)
2(4)
4(8)
7(12)
3(3)*
3(3)
39 (54)
Flore Champ.
Icon. Cent.
Afriquef
8
2
1
1
1
1
2
20
16
6
2
60
East Africa
(Pegler 1983)
8
2
3
6
1
1
7
3
5
5
7
48
Sri Lanka
(Pegler 1983)
13
5
6
2
-6
-4
1
6
43
Lesser Antilles
(Pegler 1983)
16
_
3
4
-
1
3
1
10
-3
41
Figures in parentheses = number of collections.
"Indicates collections by Henning (1895 et subseq.) of undocumented Zenker exsiccata in E.
|See Heinemann (1970 et subseq.); Beeli (1936).
species close to L. sistrata (Fr.) Quel. with a veil composed of spherocytes, but
slightly smaller basidiospores (4.4-4.8 x 2.6 /mi) and rather pinker colours. There
are also two yellow species; one is close to L. flava Beeli but with basidiospores with
a germ-pore and the second has a bright yellow citron centre to the pileus, a pale
yellow ring, and elongate-fusoid basidiospores. Eleven further collections are as yet
unplaced, five in Lepiota Sect.
Ovisporae.
Two of these represent the same species,
resembling
L.
cristata
(Fr.) Kummer
(Sect.
Stenosporae) except for the
less
projectile-
like basidiospores. Four further collections, two representing a taxon close to
L.
submontagnei
Heinem., are referrable to the genus
Leucocoprinus.
These examples
indicate how rich and underworked even this small area of West Central Africa
really is, and some collections may indeed support the suggestion that the flora there
is unique. A surprising number of the species recorded were first recognised in Sri
Lanka (Pegler 1977) and then found in East Africa (Pegler 1986). It seems as if
they will prove to be common to the Old World tropics. Some are probably pantrop-
ical having been recorded from either South America or the Southern States of the
USA or Central America, e.g.
Sericeomyces serenus
(Fr.) Heinem. and
Leucocoprinus
fragillissimus.
Probably the key to understanding the flora of lepiotoid agarics and
their significance lies with the small group, alas based on single collections, of as
yet unidentified taxa. Table 2 summarises the information available.
Conclusion
It is almost impossible to indicate the degree of endemism in the flora of larger
fungi in the Guinea-Congo Region because so little systematic collecting has been
carried out. Whereas many of the ectomycorrhizal fungi are species with a small
host range (see Buyck, Thoen & Watling this volume) probably confined to Africa,
the wood and litter decomposers include a number of species fairly widespread in
the Old World tropics, or even the tropics as a whole, in addition to African species.
The African species seem to be generally widespread south of the Sahara.
Saprophytic
fungi 345
In common with other tropical rain forests the lowland forests fringing the Bight
of Biafra extending into the Congo and south to Gabon and north to Senegal offer
a full spectrum of habitats exploited by the macromycetes. There is a continuous
supply of substrate, viz. leaf-litter, albeit often charged with phenolics and tannin
complexes, abundant twiggy debris, branches and limbs which fall through weakness,
and prostrate forest giants which when they fall through storm damage bring down
with them huge amounts of other vegetable material ranging from adjacent trees
to lianas.
The polypores and Lentinus spp. and the xylariaceous fungi generally rot the
woody material, whereas resupinates and
Marasmius
spp. and related agarics attack
the leaves. The detritus which is left is colonised by Agaricus spp. and lepiotoid
fungi. The microphyllous element of the leaf-fall is incorporated into the soil in the
Korup rain forest in about three months during the first stages of the rainy season.
A network of mycelium and rhizomorphs and fungal cords interconnect the organic
material lying on the soil surface. The only parallel found in European woodlands
is where
Hymenochaete corrugata
is found welding twigs of
Corylus
together in dense
neglected coppice (Rayner, personal communication) in parallel to
Hydrnophlebia
chrysorhiza
in Korup.
The litter decomposers may compete directly with ectomycorrhizal fungi for
available nutrients and this needs to be explored further.
The scene is now set to conduct some quantitative studies, and it is hoped future
plans will result in understanding the part played in the nutrient cycle by many of
the taxa described above.
Acknowledgements
R. Watling is grateful to his joint authors and to Kurt Hjortstam for identifying
many of the resupinate specimens, to Andrew Allo and Steve Gartlan for their help
in Cameroon and to his companions in the field, Ian Alexander, Dept of Plant &
Soil Sciences, University of Aberdeen and Jo Rother now at the National Resources
Institute, Maritime, Chatham, Laboratory help in Edinburgh has been given by
Evelyn Turnbull and the Royal Society of Edinburgh gave a Travel Grant.
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