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1 Centraalbureau voor Schimmelcultures, Fungal Biodiversity Centre, P. O.
Box 85167, 3508 AD, Utrecht, The Netherlands
2 Department of Microbiology and Plant Pathology and Department of Genetics,
Forestry and Agricultural Biotechnology Institute, Faculty of Natural and
Agricultural Sciences, University of Pretoria, Pretoria, South
Africa
3 Biosystematics Division, ARC Plant Protection Research Institute, P. Bag
X134, Queenswood, Pretoria 0121, South Africa
4 ARC Plant Protection Research Institute, P. Bag X5017, Stellenbosch, 7599,
South Africa
5 SANBI KwaZulu-Natal Herbarium, P. O. Box 52099, Berea Road, 4007 South
Africa
6 The Yellow House, Calle Aguila 12, Colonia La Maliciosa, Mataelpino,
ES-28492 Madrid, Spain
*
Correspondence: Pedro W. Crous,
crous{at}cbs.knaw.nl
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Abstract |
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 TOP Abstract INTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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Keywords Biodiversity / conservation / National Collection of Fungi / numbers of fungi / undescribed species
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INTRODUCTION |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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It is widely recognised that fungi have been relatively poorly collected and studied from most countries, regions and habitats. This is at least in comparison to plants and larger animals that are considerably easier to collect and identify than fungi. One might then have expected that the predicted numbers of fungi on earth would have been considerably greater than the 1.5 M suggested by Hawksworth (1991). Clearly different authors that have considered the likely total number of fungi have had differing views of an appropriate answer, but the discrepancy between the results of most of these studies is not particularly great. It is currently accepted that the 1.5 M estimate is highly conservative. The 100 000 that have thus far been described, therefore represents no more than 7 % of the estimated total.
One hundred years of mycology in South Africa as celebrated in this volume is not a particularly long history for mycology as science. The country is widely recognised as one of the world's biodiversity "hotspots", including areas such as the Cape Floral Kingdom, which is the smallest and most diverse biome presently known. This paper aims to summarize the major mycological developments that have happened in South Africa, as well as to provide an estimate of the number of fungi in this country. In a paper such as this it is never possible to comment on all activities and groups that have been active over the past 100 years, and for many of these there will only be a brief mention, chiefly because much of the information was not available to us at the time this paper was written. Nevertheless, it is hoped that this summary and estimate will not only be interesting, but also provide a foundation to promote and guide future mycological activity in South Africa and elsewhere.
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UNIQUE SOUTHERN AFRICAN FLORA |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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Naturally, these plants are not evenly distributed over southern Africa, and there have been various studies of the vegetation of southern Africa from the historically classic studies of Drège (1843) and Bolus (1886), through Acocks (1953 and still the most often-cited work in South African botany) to the recent studies of Low & Rebelo (1996) and Rutherford & Mucina (in prep.). For the purposes of this contribution, the concepts of biomes and centres of diversity given by Van Wyk & Smith (2001) will be used (Figs 1, 2).
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Succulent Karoo
This is the arid area to the north and west of the Cape fynbos. Different
workers have varying concepts of the boundaries of the area, giving rise to
varying estimates of the number of species present. A current SANBI project to
prepare a flora of the area is working on some 4 000 species (Paterson-Jones,
pers. comm.), but other estimates with wider geographical limits put the plant
diversity as high as 5 000 species. Many of the plants here are succulents,
but another significant part of the flora is made up of ephemeral annuals. In
both groups, it is known that not all seeds from a given year will germinate
together; up to half the crop will remain dormant until at least the second
rainy season after they have ripened.
Cape Floristic Region
The fynbos of the Western Cape has been studied for longer than any other
part of sub-Saharan Africa, and so should be among the best-known floras in
Africa. However, it is also among the world's most diverse. A recent synopsis
of the flora of this region (Goldblatt
& Manning 2000) lists some 9000 species for this area,
approximately 80 % of which are endemic. Sadly much of the area has been
replaced by, or is under threat from agriculture and urbanisation. Presently
less than 5 % of the West Coast Renosterveld is still in pristine condition.
The flora here is remarkable for reaching its greatest diversity on
essentially nutrient-free soils.
Griqualand West Centre
This region occupies the dry area bounded roughly by a line connecting the
towns of Prieska, Vryburg, Vorstershoop and Upington in the Northern Cape
Province. The vegetation is mainly various kinds of bushveld, grassland and
Karoo. There are many succulents endemic here, but the flora is not as
well-known as one may suppose. The only checklist of the plants of the area is
by Wilman (1946). Acocks
(1953) recorded over 300
species from a single sampling point in the Asbestos Hills – the highest
number from any of the thousands of points he sampled for this work.
Albany Centre
This area of endemism stretches westwards from the Kei River roughly to
Middelburg, Aberdeen and the Baviaanskloof Mountains in the Eastern Cape. The
climate is transitional between the summer-rainfall regime of the eastern part
of southern Africa and the winter-rainfall climate of the Western Cape, and at
first glance the vegetation seems transitional, too. The vegetation is highly
varied, with as many as a third (21 of 70) of Acocks's
(1953) veld types and five of
White's (1983) main
phytochoria being represented here. A flora of the Eastern Cape is in
preparation, and is expected to treat some 8400 species (Bredenkamp, pers.
comm.).
Drakensberg Alpine Centre
The highest mountain area in southern Africa includes most of Lesotho, a
very mountainous area around Barkly East in the Eastern Cape, and a fringing
area stretching along the eastern (seaward) slope of the Drakensberg from
there through KwaZulu-Natal to the mountainous area southwest of Harrismith in
the Free State. Although this area does not have an up-to-date flora, it is
well supplied with comparatively recent field guides
(Hilliard & Burtt 1987,
Killick 1990,
Pooley 2003). The last-cited
guide, estimates the flora of this centre as comprising some 2200 species, 20
% of which are endemic. Van Rooy
(2000) considers this to be
one of the main centres of diversity of mosses in southern Africa.
Soutpansberg Centre
The mountains separating the Limpopo Valley from the rest of South Africa
have long been known in botanical folklore as interesting, but have only
recently been defined as a centre of endemism, running from the Blouberg in
the west, along the Soutpansberg and including parts of the Limpopo valley
into the north-western corner of the Kruger National Park and a tiny area of
Zimbabwe. The area contains representatives of 41 % of all plant genera and 68
% of all plant families in the Flora of southern Africa area (South
Africa, Botswana, Lesotho, Swaziland, Namibia), and is certainly a meeting
place between the floras of east and west south-tropical Africa
(Hahn 1998). The only recent
account of the flora of this area specifically is Hahn's
(1994) tree checklist, but
the number of plant taxa here is estimated at about 3000.
Wolkberg Centre
Stretching from near Carolina, Mpumalanga, in the south to Haenertsburg,
Limpopo Province in the north, with a westward extension to Zebediela south of
Polokwane, this centre comprises the escarpment areas once known as the
Transvaal Drakensberg. The flora numbers between 1500 and 2000 species, some
10 % of which are endemic.
Sekhukhuneland Centre
Inland of the Wolkberg Centre escarpment is a much drier area, underlain by
basic and ultramafic rocks of the Bushveld Igneous Complex; as with the
Barberton centre below, this geology may lead one to expect many endemic
species. The flora of the area is severely undercollected, but is believed to
number some 2000 species (Retief et
al. 2001). The importance of the area as a centre of endemism
was demonstrated by Siebert
(1998), who has also done a
still-unpublished phytosociological study of this Centre
(Siebert 2001). The
vegetation is conventionally considered to be Mixed Bushveld (e.g.
Low & Rebelo 1996), but
grassland and forest remnants are also to be found there. The flora of the
Sekhukhuneland bushveld is unique. However, most of the area has been severely
degraded by creeping urbanization and subsistence farming.
Barberton Centre
The mountains between Barberton, Mpumalanga Province, and the Swaziland
border (stretching into a small part of north-western Swaziland) have a
remarkable flora with many endemics. These mountains are geologically complex,
and include some of the oldest fossil-bearing rocks on earth
(MacRae 1999), as well as
serpentinite (noted wherever it occurs for supporting unusual endemic species)
and ancient volcanic rocks. Balkwill & Balkwill
(1999) report 30 endemic
species here, many of them as yet undescribed. However, the rugged terrain
means that much of the area is effectively unexplored, and so many other taxa
of limited occurrence are no doubt still awaiting discovery
(Hurter & Van Wyk
2004).
Maputaland-Pondoland Region
This includes almost the whole of the KwaZulu-Natal Province, and a smaller
or larger fringe in surrounding areas, most notably southern Mozambique and
Eastern Cape (former Transkei). A newly-started project to generate an
electronic Flora of KwaZulu-Natal is working on a checklist of some 6500 taxa,
suggesting a flora for the whole area of about 7500 species. Vegetationally,
the area may be divided into numerous grassland, thicket and forest types.
Among the earliest specimens from KwaZulu-Natal are a series collected by
Gerrard & McKen in 1860, but serious study of the flora of the area may be
said to have started with Medley Wood, who founded the first herbarium in the
then colony in 1882 (Schrire
1983), and also collected and wrote on some fungi, mostly
associated with plant diseases. With the immense variety of plants and
habitats in this area, it is hardly surprising that undescribed fungi are
found here even more frequently than undescribed plants.
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BEGINNING OF MYCOLOGY IN SOUTHERN AFRICA |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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Most mycological activity in the post-Doidge era focused on reports of fungi causing plant diseases, with some attention to saprobic fungi, and those found to be mycotoxigenic. Reference works such as the Gorter bulletins (1977, 1979, 1981, 1982), Van der Westhuizen & Eicker (1994), and Eicker & Baxter (1999) provide lists of various groups of fungi compiled after Doidge et al. (1953). Information on fungi subsequent to 1999 can be obtained via the Internet-based electronic system of CAB abstracts (CAB) (www.cabi-publishing.org). Several lists of plant diseases caused by fungi, bacteria and viruses in South Afirica have been published. The first of these was Doidge (1924) followed by Doidge & Bottomley (1931) and Doidge et al. (1953), which was chiefly based on Doidge (1950). These records were updated in a series of bulletins published by Gorter (1977, 1979, 1981, 1982). Later, Crous et al. (2000b) published the compilation "Phytopathogenic Fungi of South Africa", which was made available online by the Systematic Botany & Mycology Laboratory in the U.S.A., and is searchable via <http://nt.ars-grin.gov/fungaldatabases/southafrica/>.
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INTENSIVELY STUDIED GROUPS |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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Subsequent to the late 1990's (Eicker & Baxter 1999), the relative dormancy in activities on ascomycetous fungi following Doidge (1950) became broken by a steady flow of contributions by plant- and forest pathologists (Agricultural News 1990), with the genera Ophiostoma Syd. & P. Syd., Ceratocystis Ellis & Halst. (Marais & Wingfield 2001, Roux et al. 2001a) and Mycosphaerella (Crous 1998, Crous & Braun 2003, Crous et al. 2004a, b) receiving particular attention. Plant-pathogenic ascomycetes occurring on Proteaceae and Restionaceae also received considerable attention, with the description of numerous new species and genera (Taylor & Crous 2000, Lee & Crous 2003a, c, d, Lee et al. 2003, 2004a). Saprobic ascomycetes occurring on Proteaceae litter, however, were less intensively studied (Lee & Crous 2003a, b, Lee et al. 2005). Contributions from other South African scientists included L. Korsten, F. Wehner, N. McClaren, B. Flett, N. Labuschagne and G. Thompson, who added valuable information pertaining to distribution records, host preferences and new disease reports.
Some ascomycete genera have received considerably more attention than others. One such genus is Mycosphaerella. Doidge (1950) listed 21 species of Mycosphaerella, and more than 100 cercosporoid anamorph species. A revision of species in this complex commenced in the early 90's, with the description of numerous new taxa from indigenous and exotic hosts (Crous & Braun 1994, Morris & Crous 1994, Crous & Braun 1995). The cercosporoids occurring in South Africa were treated by Crous & Braun (1996a), who listed 159 species from diverse hosts. Taxa occurring on Proteaceae were treated by Taylor and co-workers in a series of papers focusing on Mycosphaerella, and anamorph-genera such as Phaeophleospora Rangel, and Batcheloromyces Marasas, P.S. van Wyk & Knox-Dav. (Taylor & Crous 1999, Taylor et al. 1999). Other than Proteaceae, the Myrtaceae has also received some attention. Species of Mycosphaerella occurring on Eucalyptus were treated in several papers by Crous and co-workers (Crous & Wingfield 1996, Crous 1998, Crous et al. 2004a, Hunter et al. 2004a, b), listing 17 odd species on Eucalyptus in South Africa. The genus Syzygium, of which S. cordatum is indigenous to South Africa, was also investigated, revealing four species of Mycosphaerella on this host (Crous & Sutton 1997, Sutton & Crous 1997, Crous 1999). Given the exceptional Mycosphaerella species richness on hosts in the Proteaceae and Myrtaceae, it can be assumed that numerous species await description once leaves of other hosts are studied in more detail.
The genus Botryosphaeria and associated anamorph-genera have received considerable attention in recent years (Denman et al. 2000). Specific papers have addressed species occurring on hosts such as Eucalyptus (Smith et al. 2001, Slippers et al. 2004a,b,c,d), Proteaceae (Denman et al. 2003), Vitis vinifera (Van Niekerk et al. 2004a), Syzygium (Pavlic et al. 2004), Southern Hemisphere conifers (such as Widdringtonia) (Slippers et al. 2005b), various fruit trees (Slippers et al. 2006) and Mangifera indica (Slippers et al. 2005a), to name but a few. These studies once again indicate that there are numerous unknown species of the Botryosphaeriaceae in the Southern Hemisphere, and specifically in South Africa, awaiting description.
A suite of recent studies have focused on species that have traditionally been treated in the genus Cryphonectria (Sacc.) Sacc. & D. Sacc. (Myburg et al. 2004a, b, Gryzenhout et al. 2004, 2005a, b, 2006 – this volume). These investigations arose from the first discovery of the serious Eucalyptus stem pathogen Cryphonectria cubensis (Bruner) Hodges in South Africa (Wingfield et al. 1989). The various studies, strongly supported by DNA sequence comparisons, have shown that Cryphonectria is a genus restricted to the Northern Hemisphere (Myburg et al. 2004a, b). Related fungi occurring in Southern Hemisphere countries including South Africa, reside in various genera such as Chrysoporthe Gryzenh. & M.J. Wingf. (Gryzenhout et al. 2004), Microthia Gryzenh. & M.J. Wingf., Holocryphia Gryzenh. & M.J. Wingf. (Gryzenhout et al. 2006), Rostraureum Gryzenh. & M.J. Wingf. (Gryzenhout et al. 2005a) and Amphilogia Gryzenh., Glen & M.J. Wingf. (Gryzenhout et al. 2005b). New species and genera have also been discovered that are closely related to Cryphonectria, such as Celoporthe dispersa Nakab., Gryzenh., J. Roux and M. J. Wingf. (Nakabonge et al. 2006 – this volume).
Hyphomycetous fungi: A cursory look through Doidge (1950) reveals that none of the earlier collectors in South Africa took particular note of hyphomycetes. Of the total of 4748 species that she listed, a mere 18.5 % represented asexual forms. Subsequent studies comprised comprehensive inventories of the genera Fusarium Link (Marasas et al. 1988), Penicillium Link (Schutte 1992), Aspergillus Link (Schutte 1994), entomophagous fungi (Rong & Grobbelaar 1998), dematiaceous fungi (Sinclair & Eicker 1985, Sinclair et al. 1983, 1985, 1987, 1990, 1994), and nematode-trapping fungi (Gorter 1993b). A review of Cercospora Fresen. and similar fungi (Crous & Braun 1995), revision of the genus Cylindrocladium Morgan and related genera (Crous & Wingfield 1993, 1994, Crous 2002), and differentiation of species of Bipolaris Shoemaker, Exserohilum K.J. Leonard & Suggs and Curvularia Boedijn (Rong 2002) was compiled mostly from specimens in PREM.
The bulk of subsequent studies are miscellaneous reports of fungi from crops and animal feeds such as lucerne (Thompson 1985), Cenchrus ciliaris pastures (Bezuidenhout 1977), natural Karoo pastures (Roux 1985), as well as toxigenic representatives of Fusarium (Kellerman et al. 1972, Marasas et al. 1976) and Pithomyces Berk. & Broome (Marasas & Schumann 1972, Kellerman et al. 1980), and various synnematous and other hyphomycetes (Rong & Botha 1993, Roux et al. 1995, Jacobs et al. 2001). Other genera that received attention include Graphium Corda (Jacobs et al. 2003b), Leptographium Lagerb. & Melin (Wingfield & Marasas 1980, 1983, Wingfield 1985, Zhou et al. 2001), Cladosporium Link (Braun et al. 2003, Surridge et al. 2003), Phialocephala W.B. Kendr. (Jacobs et al. 2003a), and cercosporoid fungi (Crous & Braun 1996a, b, Pretorius et al. 2003). Several unique hyphomycetes were also reported from Proteaceae and Restionaceae in the fynbos (Wingfield et al. 1988, Mel'nik et al. 2004, Lee et al. 2004b).
Specific mention should be made of the work of Marasas on the taxonomy of the toxigenicity and phytopathologically important genus Fusarium, including soil surveys (Rheeder & Marasas 1998), and the description of numerous new species such as Fusarium andiyazi Marasas, Rheeder, Lampr., K.A. Zeller & J.F. Leslie (Marasas et al. 2001), and F. thapsinum Klittich, J.F. Leslie, P.E. Nelson & Marasas (Klittich et al. 1997). Similarly studies on the mycota of proteaceous fungi by Crous and co-workers (Crous et al. 2004a) have been very extensive.
Eyespot disease of wheat received considerable attention, with the causal organism originally being ascribed to Cercosporella Sacc., then Pseudocercosporella Deighton, and Ramulispora Miura (Robbertse et al. 1995). Crous et al. (2003) erected the genus Helgardia Crous & W. Gams to accommodate these cercosporoid anamorphs, while their discomycete teleomorphs were placed in the genus Oculimacula Crous & W. Gams. Schroers et al. (2005) recently characterised the hyphomycetes associated with guava wilt, and identified the causal organism as a species of Nalanthamala Subram.
Hyphomycetes from Vitis vinifera have been intensively studied. Halleen et al. (2004) treated the Cylindrocarpon Wollenw. complex associated with black foot disease, and introduced a new genus, Campylocarpon Halleen, Schroers & Crous. Crous et al. (1996a) erected the genus Phaeoacremonium for species associated with grapevine decline disease of grapevines and human infections, while Crous & Gams (2000) described the genus Phaeomoniella Crous & W. Gams as the main causal organism of Petri disease.
Species of Cylindrocladium Morgan (teleomorph: Calonectria De Not.) (Hypocreales) are common in tropical and subtropical regions of the world, and cause disease problems on a wide range of hosts. Schoch et al. (2000) placed teleomorphs of Cylindrocladiella Boesew. in Nectricladiella Crous & C.L. Schoch. In subsequent years Schoch & Crous (1999) reported Cylindrocladium spathiphylli Schoult., El-Gholl & Alfieri as causing Cylindrocladium root and petiole rot of Spathiphyllum, while Schoch et al. (1999) resolved the Cylindrocladium candelabrum Viégas species complex, describing the common soil-inhabiting species in South Africa as C. pauciramosum C.L. Schoch & Crous. In a recent monograph of Cylindrocladium and allied genera, Crous (2002) reported six Cylindrocladium species and five Cylindrocladiella species from South Africa.
Coelomycetous fungi: In the Doidge era, investigators often recorded coelomycetes only incidentally, usually alongside their sexual state. Because they were mainly found on plant-pathological specimens, those that were most often reported were commonly occurring members of the genera Colletotrichum Corda, Phyllosticta Pers. and Pestalotia De Not. (Doidge 1950). At this time there was a tendency to name taxa as new if they were found on new substrates. Many of these names would not hold up when judged with robust systematic techniques currently applied.
Many coelomycete reports were done from animal feed implicated in poisonings or other maladies in farm animals i.e. Phomopsis leptostromiformis (J.G. Kühn) Bubák (Van Warmelo et al. 1970) on lupins, Tiarosporella Höhn. (Sutton & Marasas 1976), Colletotrichum (Baxter et al. 1983, 1993), Urohendersonia platensis Speg. (Roux & Van Warmelo 1989), and Bartalinia robillardoides Tassi (Roux & Van Warmelo 1990).
Paraconiothyrium minitans (W.A. Campb.) Verkley was investigated as an antagonist of the devastating pathogen, Sclerotinia sclerotiorum. (Lib.) de Bary (Phillips 1985). Possibly the most notorious coelomycetes in South Africa are Phyllosticta citricarpa (McAlpine) Aa, the cause of black spot of oranges (Meyer et al. 2001), and Stenocarpella maydis (Berk.) B. Sutton (syn. Diplodia zeae van der Bijl) causing black rot of maize and intoxication of sheep (Rheeder et al. 1990). Other genera that received attention include Coniella Höhn. and Pilidiella Petr. & Syd. (Van Niekerk et al. 2004b), Colletotrichum (Lubbe et al. 2004), Harknessia Cooke (Crous et al. 1993, Lee et al. 2004a), Dothistroma Hulbary (Barnes et al. 2004), the Fusicoccum/Diplodia anamorphs of Botryosphaeria (Denman et al. 2000, Slippers et al. 2004a,b,c,d), and Phomopsis (Sacc.) Bubák (Smit et al. 1989a, b, 1996a, b, Mostert et al. 2001, Van Niekerk et al. 2005). Coelomycetous fungi occurring in the fynbos also received some attention, namely those occurring on Proteaceae (Crous et al. 2004a) and Restionaceae (Lee & Crous 2003b).
Basidiomycetes
In early years, basidiomycetes such as the Hymenomycetes
(McNabb & Talbot 1973,
Talbot 1951a,
b,
1954,
1958,
1965) and
Ustilaginales (Doidge
1950) received some attention. Approximately 36 % of the fungi
listed by Doidge (1950) are
basidiomycetous. Eicker & Baxter
(1999) present a good overview
of work done on basidiomycetes from 1977 to 1999. Their publication provides
references to studies on the genera Phaeolus (Pat.) Pat.,
Pisolithus Alb. & Schwein., Termitomyces R. Heim,
Amanita Pers., Chlorophyllum Massee, Clathrus P.
Micheli ex L., Hymenagaricus Heinem., Lepiota (Pers.)
Gray, Macrolepiota Singer, Leucoagaricus Locq. ex
Singer, Leucocoprinus Pat., Montagnea Fr. and
Hymenochaete Lév. Several new genera of heterobasidiomycetous
yeasts were also newly described through the years by J.P. van der Walt and
colleagues.
Some of the more extensive investigations were the monographic work on resupinate and stereoid Hymenomycetes and a series of papers between 1951 to 1958, dealing with Stereum Pers., Lopharia Kalchbr. & MacOwan, Cymatoderma Jungh. and the Thelephoraceae (Gorter 1979). Early investigations also included a series of papers on tree pathogens and wood-destroying Hymenomycetes (Van der Westhuizen 1972).
Some revisionary work has been done on collections of certain taxa at PREM in the course of monographic work at the University of Buenos Aires, Argentina, namely species of Hymenochaete Lév. (Hymenochaetaceae) (Job 1987). Limited information is available about the Geasteraceae (Coetzee & Eicker 1994, Coetzee & van Wyk 2003) but a revision of the South African Stereum species, and the Geasteraceae (earth stars), which will to a large part rely on material lodged in PREM, is in progress (J. Coetzee, pers. comm.).
Significant contributions following Doidge were made by D.A. Reid from the Royal Botanic Gardens Herbarium, Kew. Reid reappraised the type and authentic specimens of species of Basidiomycota described from South Africa in PREM (Van der Westhuizen & Eicker 1994), the van der Bijl Herbarium (now housed at PREM), and elsewhere (Reid 1975). This has been the foundation for his documentation of South African mushrooms in collaboration with Albert Eicker from the University of Pretoria. An equally productive partnership has provided us with a scientific guide to our edible and poisonous mushrooms and other large fungi. In 1994, G.C.A. van der Westhuizen's lifetime of mycological research and photography culminated in a field guide (Van der Westhuizen & Eicker 1994), with excellent colour photographs of some 160 species of local macrofungi. The most recent studies include DNA phylogenetic data, like that published on species of Termitomyces R. Heim (Botha & Eicker 1991a, b, De Fine Licht et al. 2005).
Studies during the early part of the last Century reported Armillaria mellea (Vahl: Fr.) P. Kumm. in South Africa (Pole Evans 1933, Kotzé 1935, Bottomley 1937). These were largely associated with an expanding plantation forestry industry and the fact that this fungus resulted in tree death. A series of recent studies have shown that the fungus killing trees in this country is A. fuscipes Petch (Coetzee et al. 2000) and that there are probably at least two other species occurring in neighbouring countries such as Zimbabwe (Mwenje et al. 2003). Intriguingly, it has also been shown that the Northern Hemisphere species, A. mellea was introduced into South Africa, probably by the early Dutch settlers (Coetzee et al. 2001). Likewise, the Northern Hemisphere species A. gallica Marxm. & Romagn. has recently been recorded from dying Protea plants in the Kirstenbosch botanical gardens (Coetzee et al. 2003).
G.L.I. Zundel of the U.S.A., who collaborated with Pole Evans and Doidge in describing and re-investigating material found by South African mycologists, extensively studied local smut fungi. K. Vánky in Germany radically changed the taxonomic study of this group of fungi by advocating the use of morphological characters obtained by germinating spores (Vánky 1997, 1999a, b, 2000a–b, 2001). It is problematic when we apply these methods to old herbarium specimens, however, because their spores have lost the ability to germinate. One of our earliest records and most striking smuts is maize boil smut, Ustilago maydis (DC.) Corda [= U. zeae (Link) Unger].
Rust fungi (Uredinales): Most of the rust fungi known from southern Africa were treated and described by Doidge (1927, 1928, 1939, 1940, 1948a, b). This suite of papers remain the basis for identification of these fungi in southern Africa, and are relevant to the whole of the African continent. All known species were listed in Doidge (1950). A total of 474 species are listed from southern Africa, including 145 anamorphs.
Since Doidge (1950), nine species have been transferred to other genera, and a further 30 species names have been reduced to synonymy. In addition, eight anamorph species of Aecidium Pers. have been connected to their teleomorphs (Kleinjan et al. 2004), and three have been demonstrated to be endocyclic (Wood 1998, 2004, Wood & Crous 2005), whilst 11 anamorph species of Uredo Pers. have been connected to their teleomorphs. Many of these changes were published in Jørstad (1956) and various publications summarised in Cummins (1971). It can be expected that the status of many more names will change as further taxonomic studies progress (Van Reenen 1995).
Since Doidge (1950), a number of new species have been described by G. B. Cummins and H. Gjærum in various papers, none dealing exclusively with species from the region (Cummins 1960, Gjærum & Reid 1983, 1998, Gjærum 1988a, b, 1999). More new species have been described recently (Shivas 1991, Wood 2002, Mennicken et al. 2003, Mennicken & Oberwinkler 2004, Mennicken et al. 2005a, b, Wood & Crous 2005, Wood & Scholler 2005). Taking these above-mentioned changes into account, there are currently 537 species of rust fungi, representing 40 genera and 10 families presently recorded from southern Africa (A. Wood, unpubl. data). This tally will increase, as several more new species and records await publication. Of the countries included, southern Angola, Botswana, Mozambique, Namibia and Swaziland have few species recorded, and Lesotho has none. Only South Africa and Zimbabwe are relatively well explored mycologically, though little or no collecting has been done in large areas of even these two countries. A total of 20 new species have been described from the Western and Northern Cape Provinces of South Africa, and Namibia, in just the last 5 years (Wood 2002, Mennicken et al. 2003, Mennicken & Oberwinkler 2004, Mennicken et al. 2005a, b, Wood & Crous 2005, Wood & Scholler 2005), demonstrating that many new species await discovery in the subregion where there has been little or no collecting in the past. However, the number of species known to occur is probably not a gross underestimation of the actual total present, as with the study of fresh specimens it is probable that a number of species will be reduced to synonyms, and many anamorphic species will be linked to their teleomorphs.
Hennen & McCain (1993) suggested that the number of rust species in any area would be at least 5 % of the number of vascular plants. In South Africa and Zimbabwe the figures are only 1.7 and 3.2 %, respectively (Table 1). As stated above, the number of species is not considered likely to rise considerably. Why then are there relatively few rust species recorded from the subcontinent in comparison to other countries (Table 1)? There are a number of possible explanations for this, the most likely including: (1) Much of South Africa, as well as Botswana and Namibia is semi-arid to desert. The low rainfall associated with these areas would restrict opportunities available to rust fungi to infect their hosts, which could result in the lower diversity of rust fungi. (2) There are very few rust species recorded from the fynbos and succulent Karoo biomes, despite the high plant diversity for which these biomes are well known. The high plant species diversity is largely produced by few but speciose genera and families with high turnover of species over short distances (Linder et al. 1992, Goldblatt 1997).
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The rust fungi in South Africa tend to either occur on one or a few closely related plant species with a large distribution, or on numerous related plant species, many of which have limited distributions. Endophyllum osteospermi (Doidge) A.R. Wood, E. elytropappi (Henn.) A.R. Wood & Crous (Wood & Crous 2005) and Uromyces kentaniensis Doidge are examples of the former, whilst U. bolusii Massee, U. ixiae (Lév.) G. Winter and Puccinia byliana Dippen. are examples of the latter (A. Wood, unpubl. data). The greatest diversity of rust fungi in South Africa is in the eastern parts, associated with a greater amount of summer rain. There the ratio of rust fungi to plant species is probably approximately the same as for Zimbabwe, which is ecologically similar.
Zygomycota
Only seven genera of the Mucorales were noted by Doidge
(1950), one of which,
Haplosporangium Thaxt., has not since been found. They include the
important saprotrophic species, identified early on, such as the ubiquitous
Rhizopus stolonifer (Ehrenb.) Vuill. – listed as R.
nigricans Ehrenb. – which causes post-harvest decay, particularly
in sub-tropical fruit. A subsequently recorded species, R. oryzae
Went & Prins. Geerl., has been associated with mycoses in humans. It is
the most commonly found member of this group, particularly in fodder samples.
During the course of several years, PREM has supplied numerous cultures of
Mucorales, and participated in research focused on metabolite
studies. Certain strains produce gamma linoleic acids
(Botha et al. 1995)
culminating in patenting of a fermentation process.
By 1999 the number of genera from this group known to South Africa had risen to 20 (Roux 1996). A study on Trichomycetes, parasites or commensals in the digestive tract of arthropods, yielded several new records and new species (Gorter 1993a).
Lichen-forming and lichenicolous fungi
In comparison with other fungal groups, the macrolichens of South Africa in
particular are relatively well-studied, though much needs to be done on the
microlichens and lichenicolous species. Early records were compiled in
Stizenberger
(1890–1895), but the
major synthesis to date is that of Doidge
(1950). Extensive collections
were made by Elise Esterhuysen (1912–) in the late 1940s–1950s
(material in the Bolus Herbarium, University of Cape Town), but the principal
contributions have been by Ove Almborn (1914–1992) from Lund (Sweden)
and Franklin A. Brusse (1951–) who was for some years on the staff of
the National Herbarium in Pretoria. Almborn specialized on
Teloschistes Norman (Almborn
1989) but travelled extensively in the country, for example
spending 5 mo there in 1953 (Almborn
1966), analyzing distribution patterns (Almborn
1987,
1988), and also issued an
important exsciccate, "Lichenes Africani"; this comprised six
fascicles from 1956–1991 including 150 numbers. Brusse had his major
interest in the Parmeliaceae and described many species new to
science, especially effigurate-crustose species on rock later placed in the
endemic genus Karoowia Hale (the species known from Australia appear
to belong elsewhere). Numerous other lichenologists have visited and
collected, amongst the most notable being Gunnar Degelius (1903–1993)
who specialized in Collemataceae
(Degelius 1974), Ingvar
Kärnefelt (University of Lund) with a special interest in
Teloschistaceae, Leif Tibell who specializes in caliciaceous groups
(who visited with colleagues from the University of Uppsala in 1997), and
Mason E. Hale jr (1928–1990) who described numerous species and several
new genera in the Parmeliaceae (e.g. Hale
1986,
1988), although not all his
new genera have yet been evaluated by molecular methods. Much collecting was
also carried out during the International Association for Lichenology's field
meeting in Namibia and South Africa in 1986, but the results from that
excursion have not been synthesized. Some revisionary studies have been
carried out based on the collections of these and other lichenologists, for
example the monograph of Diploschistes Norman in South Africa by
Guderley & Lumbsch
(1996).
More recently, as a part of the Biota-So5 project funded by the German government, an extensive survey of lichen biodiversity from the Cape Province to northern Namibia is underway, the leading researcher involved being Tassilo Feuerer (University of Hamburg). In addition, an international expedition led by Ana Crespo (Universidad Complutense de Madrid) and focussing on the Parmeliaceae made about 750 collections in May–June 2005 from the Western Cape north to the Namibian border.
Sadly there is no systematic lichenologist resident and active in South Africa today. Further, there is no modern critical checklist available, but a preliminary list compiled from the literature (www.biologie.unihamburg.de/checklists/africa/southafrica) contained 1716 species as of 1 March 2005. This list is a valuable basis for future work but is very preliminary and includes both many early names copied from the literature whose position and status needs to be reassessed, and others that have been revised but have yet to be updated from other literature. A reasonable estimate of the total number of lichen-forming species to be expected, by comparison with checklists from other regions of the world with similarly diverse climatic regions and rock types (Will-Wolf et al. 2004) would be about 2000 (excluding lichenicolous fungi).
In comparison to the flowering plants, only a handful of genera appear to be endemic, notably Combea De Not. and Karoowia s. str., with others requiring reassessment, for example Almbornia Esslinger, Canomaculina Elix & Hale, Namakwa Hale, and Xanthomaculina Hale. In contrast, there are many endemic species, especially in the Parmeliaceae and the Teloschistaceae. The lichen assemblages are very distinctive in different parts of the country, as discussed by Almborn (1988), comprising at least eight phytogeographic categories: ubiquitous, steppe and desert (e.g. Namaqualand, Karoo), montane (over 1000 m alt.), oceanic (e.g. Table Mountain, Drakensberg Mountains), tropical-oceanic (e.g. northern KwaZulu-Natal, Mpumalanga), maritime (on coastal rocks), and endemic.
Lichenicolous fungi, fungi obligatory on lichens as pathogens, commensals or saprobes, have hardly been reported from South Africa. None are included in the 2005 preliminary checklist, not even representatives of such ubiquitous genera as Abrothallus De Not., Lichenoconium Petr. & Syd., Polycoccum Saut. ex Körb., and Stigmidium Trevis. There are, however, some scattered records, such as that of Lichenostigma cosmopolites Hafellner & Calat. (Hafellner & Calatayud 1999) which is very common on Xanthoparmelia (Vain.) Hale species in the country (D.L. Hawksworth, pers. obs.), and an undescribed Polycoccum on Karoowia adhaerens (Nyl.) Hale (Váczi & Hawksworth 2001). Many lichenicolous fungi were collected during the visit of Ana Crespo's group in 2005, but have yet to be fully identified. In comparison with what is known of the species richness of lichenicolous fungi compared with the number of potential host lichen species, for example 403 vs. 1677 lichen species in Great Britain and Ireland (Hawksworth 2003), around 475 species would seem to be a reasonable estimate of the number to be expected in South Africa.
Oomycetes
Fifteen species of Phytophthora de Bary from 74 different hosts
have been recorded from eight southern African countries during the period
1941 to 2001. Typical disease symptoms were damping off, root rot, stem rot,
stem canker, crown rot, tip blight, leaf rot and fruit rot. Records for
Pythuim Pringsheim species date from 1931 to 2004. Twenty species of
Pythium have been reported from 65 different hosts, and
representative disease symptoms were damping off, seedling blight, root rot,
stem rot (foot rot), crown rot (heart rot), fruit rot, tuber rot and soft
rot.
Phytophthora: The first detailed study of Phytophthora species with associated hosts and distribution patterns in South Africa, was done by Wager (1935, 1941) describing P. cactorum (Lebert & Cohn) J. Schröter, P. cinnamomi Rands, P. citrophthora (R.E. Smith & E.H. Smith) Leonian, P. cryptogea Pethybridge & Lafferty, P. infestans (Montagne) de Bary, P. nicotianae Breda de Haan and P. syringae (Klebahn) Klebahn. Mes (1934) and Wijers (1937) reported P. cactorum on various hosts and included descriptions and disease symptoms. Other studies confirmed species identification (Wager 1931, van der Merwe et al. 1972). Descriptions of various Phytophthora species from different vegetable and ornamental crops were recorded by Thompson (1981), Thompson & Phillips (1988), Ferreira et al. (1991), Thompson & Naudé (1992) and Thompson et al. (1994). Morphological and molecular methods (RFLPs of total DNA) were used by Botha (1993) to confirm species identity of P. nicotianae on tree lucerne. Linde et al. (1997) applied isozymes to distinguish two separate populations of P. cinnamomi, determining genotypic variation within A1 and A2 isolates, and established that sexual reproduction was rare or absent in South African isolates. Linde et al. (1999) used both RAPDs and RFLPs to assess genotypic diversity and reveal DNA polymorphisms in South African and Australian isolates of P. cinnamomi. Von Maltitz & von Broembsen (1984) reported P. porri Foister from onions, followed by P. citricola Sawada causing shoot tip blight of lemons in propagation tunnels (Von Maltitz & von Broembsen 1985).
Comprehensive disease surveys, but with limited taxonomic treatment, were performed by Marais (1979) in the Western Cape Province, surveying grapevine rootstock diseases. Von Broembsen (1984) did an extensive survey of P. cinnamomi root rot in indigenous fynbos, and from the major catchment rivers in the Western Cape. Thompson (1987, 1988) surveyed alfalfa in various provinces for P. medicaginis E.M. Hansen & D.P. Maxwell and P. drechsleri Tucker. Linde et al. (1994a, b) surveyed the major commercial forestry areas in South Africa for root rot caused by oomycetes, while Thompson et al. (1995) surveyed citrus roots for Phytophthora and Pythium in the Mpumalanga and Limpopo Provinces. Labuschagne et al. (2000) reported P. capsici Leonian causing wilt of pumpkin, and stem and root rot of tomato (Labuschagne et al. 2003). McLeod et al. (2001) conducted an extensive survey of Late Blight (P. infestans) in the major potato production areas in South Africa, determining mating type and using various molecular markers to characterise isolates.
Pythium: The first major contribution by Wager (1931) consisted of several species descriptions and disease symptoms on hosts. Additional species descriptions, temperature-growth measurements and hosts, was also reported by Wager (1941). Doidge (1950) and Doidge et al. (1953) compiled the host range of numerous Pythium species, and this compilation was updated by Gorter (1977, 1982) and Crous et al. (2000b). Darvas et al. (1978) and Darvas (1979) added further records of Pythium species with host records. Scott et al. (1979) and Scott (1987) made a major contribution with regard to the occurrence and disease symptoms of Pythium species from various small grain crops. An extensive overview of all recorded Pythium species in South Africa from 1926 to 1989 was compiled by Denman & Knox-Davies (1992) and included distribution, host range and species diversity. A detailed taxonomic description of various Pythium species associated with vegetables was reported by Botha & Coetzer (1996) and Pythium prolatum W.A. Campbell & Hendrix causing wilt of Rhododendron sp. (Botha & Crous 1992). Pythium species have also been associated with grapevine (Marais 1979, 1980), medics (Lamprecht et al. 1988), lucerne seedlings (Denman & Knox-Davies 1992), pines and eucalypts (Linde et al. 1994a, b) and lettuce (Labuschagne et al. 2002). Reports of asexual, sporangial isolates of Pythium, viz. G-group, F-group, T-group and P-group as well as homothallic species, e.g. P. coloratum Vaartaja, P. irregulare Buisman, P. myriotylum Drechsler, P. perplexum Kouyeas & Theohari, P. dissotocum Drechsler and P. spinosum Sawada, have been reported from closed hydroponic systems in South Africa (Labuschagne et al. 2001, 2002, Gull et al. 2004, Tesfaendrias et al. 2004).
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MISCELLANEOUS REPORTS FROM LITTLE-STUDIED NICHES |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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Fungi associated with mycorrhizal associations have also received some attention in South Africa. A significant contribution was the ecological survey of arbuscular mycorrhiza made by Wehner in 1976 (Eicker & Baxter 1999). Subsequent studies include recordings of Acaulospora laevis Gerd. & Trappe, and investigations on fynbos soils by Mitchell, Reid, Straker, van Greuning, Sinclair and later during a survey of the fungi associated with Pinus by Van der Westhuizen and Eicker (Eicker & Baxter 1999).
Several papers were published on fungal endophytes. Crous et al. (1995a) listed 55 endophytes from Triticum aestivum, while Serdani et al. (1998) treated the endophytic Alternaria Nees species associated with core rot of apples. Kriel et al. (2000) provided an overview of foliar endophytes in Gymnospermae, and Mostert et al. (2000) treated the endophytic Phomopsis spp. occurring in grapevines. Smith et al. (1996) treated endophytic Botryosphaeria species in Pinus and Eucalyptus. Swart et al. (2000) reported on the fungal endophytes in cultivated Protea, Leucospermum and Leucadendron, and Taylor et al. (2001) treated the endophytes occurring in Protea spp. grown in the wild.
Several papers focused on leaf-litter fungi in general (Sinclair & Eicker 1985, Sinclair et al. 1985, 1987, Crous 1990, Sinclair et al. 1990, 1994), or on specific hosts such as Syzygium cordatum (Crous et al. 1995b), Podocarpus (Crous et al. 1996b), and Eucalyptus (Crous et al. 1990a, Crous & van der Linde 1993, Crous et al. 1997).
Fungi on commercially important exotic hosts such as Eucalyptus, Pinus and Acacia have been studied in detail. Crous et al. (1990b) listed the shoot and needle diseases occurring on Pinus spp., while the leaf pathogens occurring on Eucalyptus were treated Crous et al. (1989). Viljoen et al. (1992) provided a summary of the fungal pathogens on Pinus and Eucalyptus seedlings in nurseries in South Africa. Crous (1998) treated the species of Mycosphaerella occurring on Eucalyptus, and listed 14 species from South Africa, while Slippers et al. (2004d) treated the Botryosphaeria spp. occurring on this host. Roux & Wingfield (1997) treated the fungi causing diseases of A. mearnsii, and found various species of fungi, including Fusarium graminearum Schwabe and Ceratocystis albifundus M.J. Wingf., De Beer & M.J. Morris to be associated with a wilt disease of this host (Roux & Wingfield 1997, Roux et al. 2001a, b).
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FUNGI ON INDIGENOUS HOSTS: FABULOUS FYNBOS FUNGI AS A CASE STUDY |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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In a study aimed at exploring the saprobic (litter) microfungi inhabiting these two families, nature reserves and botanical gardens were visited over a 2-year period (2000–2001). About 580 fungal strains were isolated from 34 restio species (representing 15 genera), which consists of approx. 150 fungal genera and 180 species. Another 580 fungal strains were isolated from 43 protea species (representing five genera), which consists of approx. 120 fungal genera and 185 species. A total of about 230 fungal genera were isolated, of which 190 were confined to either the one or the other host family, while 40 fungal genera occurred on both families. A total of 380 were identified to species level, of which 355 were restricted to one or the other family, while 25 species occurred on both families (Fig. 4).
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The number of fungal species isolated from each collection varied depending on host plant habitat. For example, 38 fungal species were isolated from Brabejum stellatifolium (Proteaceae), which has a riverine habitat and thick twigs and branches. In contrast, only four species were isolated from Elegia filacea (Restionaceae), which grows in dry areas, and has culms of approx. 1 mm diam.
From these data we have found that there are at least three unique species of saprobic microfungi for each species of Proteaceae or Restionaceae thus far investigated (Fig. 5). However, we regard this as a minimal estimate because of the limitations of the damp-chamber isolation technique, and the undersampling of microhabitats such as infructescences.
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FUNGI ASSOCIATED WITH INSECTS |
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TOPAbstractINTRODUCTIONUNIQUE SOUTHERN AFRICAN FLORABEGINNING OF MYCOLOGY IN...INTENSIVELY STUDIED GROUPSMISCELLANEOUS REPORTS FROM...FUNGI ON INDIGENOUS HOSTS:...FUNGI ASSOCIATED WITH INSECTSFUNGAL ESTIMATES FOR SOUTH...TO COLLECT, STUDY AND...References |
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One group of insect-associated fungi that has been better characterised than most, especially in the Northern Hemisphere, is that of the ophiostomatoid fungi with bark and ambrosia beetles (Six 2003, Kirisits 2004). In South Africa, three Raffaelea Arx & Hennebert species have been described from two native ambrosia beetles (Scott & Du Toit 1970), while at least 11 ophiostomatoid species have been reported from introduced pine bark beetles (Zhou et al. 2001, 2006). Unfortunately the number of ambrosia and bark beetle species in South Africa has not been determined to date. Seven novel ophiostomatoid species have also been described from merely nine of the more than 300 Protea spp. in South Africa (Marais et al. 1998, Marais & Wingfield 2001, Roets et al. 2006). The morphology of the fungi and the wealth of insects present in protea infructescences suggest that the species from proteas are vectored by arthropods (Marais & Wingfield 2001). Five more, undescribed Sporothrix Hektoen & C.F. Perkins and Ophiostoma spp. were recently discovered (De Beer et al. 2005) growing on fungal combs in 13 termite mounds, representing only three of the at least 42 fungus-growing termite species (Macrotermitinae) in Southern Africa (Uys 2002). These studies on ophiostomatoid fungi associated with beetles, Protea spp. (and their insects), as well as
