Mini-Review

Where Are All the Undescribed Fungi?

David L. Hawksworth and Amy Y. Rossman

First author: International Mycological Institute, Bakeham Lane, Egham, Surrey TW20 9TY, U.K.; second author: Systematic Botany and
Mycology Laboratory, USDA Agricultural Research Service, Beltsville, MD 20705.
Accepted for publication 29 May 1997.

ABSTRACT

Hawksworth, D. L., and Rossman, A. Y. 1997. Where are all the unde- In addition to undiscovered species, many have been collected but remain
scribed fungi? Phytopathology 87:888-891. lost or hidden as named species and ignored for lack of modern char-
acterization; others have been collected and recognized as new species
The hypothesis that there are 1.5 million fungal species on Earth, of but remain undescribed. Some fungal species are unrecognized within
which only about 70,000 are described, implies that 1.43 million remain erroneously circumscribed species, often based on presumed host speci-
undescribed. The recognition that many new species have yet to be found ficity, while others exist as biological species but remain buried within
is of fundamental importance to plant pathologists, agronomists, and those broadly defined species for lack of gross morphological charac-
plant regulatory officials, among others, who continue to encounter dis- terization. From these data, one must conclude that enormous numbers of
eases caused by previously unknown or understudied fungi. Unexplored unrecognized fungi can be found almost everywhere, including one’s own
habitats with their arsenal of unknown fungi are also of interest to those backyard.
searching for novel organisms for use in biological control or for their
pharmaceutical attributes. This paper presents data on the expected num- Additional keywords: biodiversity, biosystematics, diversity, emerging dis-
bers of fungi in some relatively unexplored habitats, such as tropical eases, inventory, lichenized fungi, plant pathogens.
forests, and those obligately associated with plants, lichens, and insects.

The generally accepted estimate of the number of species of have been undertaken that provide supportive evidence (8,19). For
fungi on Earth is a conservative 1.5 million (17,19,25). This estimate example, more than 500 fungi were identified in intensive studies
was derived by extrapolating both from data on known fungi from of litter from only five tree species in Panama (11). After 1 year of
well-studied regions, as well as data on well-studied fungi from intensive collecting in one forest in Kenya, 59 of 75 (79%) of the
plant hosts (17). Estimates of the species numbers in each major leaf-dwelling ascomycetes encountered were new species (40).
fungal group corroborate this conclusion (45). Although it is un- Recent investigations of fungi on palms yielded 1,580 species, of
certain exactly how many fungal species are already known, one which 75% of the species collected were new to science (27).
can be reasonably confident that it is in the range of 72,000 (21) to Data on the primarily tropical ascomycete family Phyllachoraceae,
100,000 (45). This implies that we know as little as 5%, i.e., only tar leaf spot fungi, suggest that although 1,150 species have been
1 in 20 of the species of fungi that exist. Clearly the knowledge described, the family may actually be support 169,000 species (7).
gap is immense. But where are the remaining 1.43 million species In exploring tropical regions for fungi, it is anticipated that the
to be found? The answer to this question is essential for mycol- most widespread and common species in a site will tend to be
ogists and plant pathologists to be able to anticipate and respond found first. Most likely, such species have been collected before
to problems resulting from emerging diseases caused by previ- and are described already. However, the longer the time spent in
ously unknown or understudied species and to prioritize and direct intensive exploration, the more species are discovered, as evi-
resources toward understanding the systematics of the most im- denced by the continuing accounts of new species, primarily of
portant and unknown groups of fungal plant pathogens. Addition- mitotic fungi, that have been discovered in Taiwan during the past
ally, the answer to this question will test the hypothesis itself and, 16 years (29–37). A 2-month visit to Papua New Guinea yielded 6
thereby, address the skepticism of some nonmycologists about the genera and 89 species of lichen-forming and lichenicolous fungi
number of undescribed species of fungi (16,39). In this paper, we new to science (2). Only through an intense examination of all
draw attention to results that indicate where at least some of this fungi in a defined area will the finite number of fungal species be
enormous treasure-trove of fungi is to be found. determined. The results of a recent workshop to plan an all-taxa
biodiversity inventory in the Área de Conservación Guanacaste, Costa
TROPICAL FORESTS Rica, estimated the number of fungal species remaining to be en-
countered in the 120,000-ha reserve at around 50,000 (6).
Many biodiversity specialists, particularly those with expertise An overall indication of whether the tropics are more species-
in macroorganisms, have determined that tropical forests are more rich in fungi than temperate regions might be expected based on
species-rich than temperate forests; thus, this is generally assumed an analysis of the origin of newly described fungi. The following
to be true for fungi as well. However, this hypothesis has not been data were extracted from the Index of Fungi database maintained
rigorously tested for fungi, although a few comprehensive studies at the International Mycological Institute (IMI), Egham, U.K.
Tropical countries were the source of 49% of the 16,013 fungal
species, including fossils, described as new during a 10-year per-
Corresponding author: D. Hawksworth; E-mail address: d.hawksworth@cabi.org iod, 1981 to 1990. However, the United States was the origin of
the largest number of new taxa, with 1,623 (10.1%) species, fol-
Publication no. P-1997-0716-01V
This article is in the public domain and not copyrightable. It may be freely re-
lowed closely by India, with 1,554 (9.7%) species. Of the remain-
printed with customary crediting of the source. The American Phytopathological ing countries with more than 1% of the total, 8 of 22 were tropi-
Society, 1997. cal. Rather than indicating whether tropical countries are the richest
888 PHYTOPATHOLOGY
source of new fungal species, these data suggest that all countries trees nearby. Well-designed studies are needed to provide reliable
are inadequately known mycologically. Those yielding the most data on this important issue.
species new to science are merely the ones that receive the most Data on fungi reported on Eucalyptus recently have been com-
intense scrutiny rather than those that are inherently the most piled (46). Of the known 450 species in Eucalyptus, 1,350 species
species-rich. This view is supported by the situation in the British of fungi are known from 150 host species. The number of fungi
Isles, the most comprehensively studied region for fungi in the from each host species ranges from 1 to 282. The highest number
world. Here critical or intensive studies invariably yield species of fungi were found on E. globulus, and 150 of the 282 fungi on
that have never been described: 460 species new to science were that host were not listed on any other species of Eucalyptus. Anal-
described from Great Britain and Ireland during the same 10-year yses of these and other data sets suggest that 5.3 unique fungi per
period. host plant species is a reasonable working number (20).

UNEXPLORED HABITATS Lichens

Lichenicolous fungi, i.e., fungi growing on and in lichens, are a
The number of habitats that potentially support specialized and largely undiscovered group of organisms, many of which are nov-
unique fungi is enormous. The fungi described as new to science el genera and species. The number catalogued has increased from
during 1981 to 1990 (mentioned above) were associated with 1,982 457 species in 1976 to 894 in January 1996, a rise of 96% during
host genera or substrata. Some previously unexplored substrata 20 years. These totals are set to increase even more steeply, as
and habitats from which these fungi were found include the rumens evidenced by the following four studies. In a recent preliminary
of herbivorous mammals; algae, lichens, and mosses; marine account of heterobasidiomycetes growing on lichens, 53 species
plants, including mangroves and driftwood; rocks; and insect scales. were recognized, of which 46 (92%) were previously undescribed
Soil and plant litter also continue to be a rich source of unusual (13). An examination of fissitunicate pyrenomycetes on tropical
fungi. Selected examples from a growing number of data sets are foliicolous lichens yielded 49 species, of which 36 (73%) were
indicative of our current lack of knowledge. new to science (38). In examining fungi from the relatively well-
studied lichen Peltigera, 96 species are now known, 61 of which
Plants occur only on this lichen host; 7 of these species were described
The 200,000 species of vascular plants continue to serve as the only within the past year (22). A study of the fungi isolated from
major reservoir of novel fungi. It has been estimated that there thalli of two fruticose lichen genera from one site in Germany
may be around 270,000 species of plant-pathogenic fungi in the revealed 506 strain types (42).
tropics (47). As in the case of geographic analysis, it is intensity
of study that best explains the observations. For example, during Insects
the 70-year period from 1920 to 1990, about three times as many In preparing estimates of the global numbers of fungi, Hawksworth
new species of fungi were described from the relatively well-studied (17) adopted a conservative approach to both insect numbers and
Poaceae compared to the related but less-studied Cyperaceae (8). the extent of host specificity of the insectivorous fungi. New data
Within the Poaceae, the number of fungi per potential host species have been presented on the extent of host specificity in the insect-
ranged from a low of 0.3:1 in the genus Poa to 35:1 in the genus infecting Laboulbeniales, a group of fungi that occur on the exo-
Zea, with 387 reported on Zea mays alone. Even among grasses in skeletons of beetles and flies. Detailed comparisons were made
the United States, there are no reports of fungi on 45% of the between rich collections from sites in the United Kingdom and
grass hosts (15). After analyzing these data (15), Clay (10) re- Sulawesi in Indonesia (50). This analysis suggests that, al-
ported that among the 20 grasses with the highest reported num- though only 1,855 species of Laboulbeniales are currently
bers of fungi in the United States, only 3 are native grasses. These known, the actual number is likely to be between 10,000 and
numbers reflect the intensity of study, i.e., economic importance, 50,000 species. This implies that only 3.7 to 18.5% of the spe-
rather than the biology of the host plants or the true fungal bio- cies in the order are known. The situation may be even more
diversity of these two related plant families. extreme in the case of the Trichomycetes, a group of enigmatic
The same situation is found when one plant species becomes fungi occurring in the hindguts of insects and other inverte-
the focus of attention. In a search for potential fungal biocontrol brates. These organisms are so poorly studied in most regions
agents, the pernicious weedy shrub Lantana camara was scruti- of the world that generalizations are hazardous. However, it is
nized carefully and is now known to support 55 fungi, of which 28 clear that these are a major source of undiscovered fungi even in
appear to be restricted to that one host species (3). Three new the United Kingdom (41).
species, one representing a new genus, were discovered during
one study of L. camara in Brazil (3). The maritime rush Juncus New Techniques Applied to Previously Studied Substrata
roemerianus, found along the eastern coast of the United States, Soil is one of the substrates from which fungi have been studied
was not known to support any host-specific fungi until two re- for many years, resulting in some relatively comprehensive ac-
searchers began investigating its mycota in 1991. From this one counts of soil fungi (14,44). Yet, the results of application of se-
plant species, Kohlmeyer et al. (28) have now described 20 new lective methods for isolating fungi from soil, such as selective iso-
species, including 8 new genera and 1 new family. Even on this lation of low-temperature microfungi (9) or basidiomycetes (49),
single-host plant, each of the numerous microhabitats are occu- demonstrates that entire groups of fungi have been overlooked,
pied by different fungal species. even in relatively well-studied substrata. Species occurring at low-
In the tropics, it is particularly difficult to ascertain patterns of er frequencies and those that are technically difficult to detect are
species richness. A. C. Batista and coworkers (12) recorded 3,340 less likely to have been discovered.
fungi from Brazil associated with 523 plant species, equivalent to The increased use of techniques that favor detection of rare spe-
an average of 6.4 fungi per host plant studied. Such data were not cies, specifically particle filtration, has resulted in the isolation of
collected for an analysis of species richness and are difficult to several hundred fungi from one substrate without reaching an
assess, because not all plants were studied and plant species with- asymptote in the species-discovery curve (4). This technique, in
out fungi were not listed. Further, although several new species which washed, pulverized, and filtered substrates are placed on a
could occur on a single leaf, no information is available from weak agar medium and closely observed, allows elimination of
which to make sound extrapolations. How many leaves on the one fast-growing fungi, so slower growing species can be detected.
tree species had novel fungi was not specifically reported nor was Using this technique, the number of fungal species isolated from
how many of those fungi occurred on different and even unrelated the decaying plant parts of a single tropical plant exceeded 200
Vol. 87, No. 9, 1997 889
(4,43). In comparing species diversity from one plant host deter- Collected and Unidentified
mined by two detection methods, the particle filtration technique Most mycologists concentrate on one or two major groups of
yielded 10 times the number of fungal species detected by moist- fungi that they collect and study throughout their lifetime. With
chamber incubation (43). Thus, methods of isolation and observa- the opportunity to visit previously unexplored regions of the world,
tion play an important role in determining fungal diversity. each scientist bears the responsibility of collecting specimens of
all groups of fungi and distributing them to experts. Yet, for many
LOST OR HIDDEN SPECIES groups of fungi, including plant pathogens, no expert exists to
study the specimens. At best, the specimens are deposited in the
Many fungi are lost or hidden within existing constructs. world’s herbaria and identified only to the genus or not at all.
Cryptic Biological Species Almost every mycological collection has folders, drawers, or boxes
of specimens that represent a bonanza of as-yet unstudied and,
Fungi traditionally have been distinguished on the basis of mor- possibly, undescribed species. Experts are needed who can under-
phological characters. However, genetic and molecular data sug- take the rigorous study required to provide a formal description
gest there are many functional biological species masquerading and place each new species in its taxonomic framework. In the
under a single species name or recognized only at an infraspecific case of major mycological institutions, these may amount to sev-
rank or as a “special form.” It is striking that almost every fungus eral thousand specimens and represent a significant portion of the
critically studied by population biologists is found to comprise a undiscovered fungal biodiversity. To speculate, the number of un-
number of reproductively isolated “biological species” (5). In dis- described fungal species represented by these specimens is prob-
tinguishing such biological species as formally described taxa, the ably more than 20,000.
biological information characteristics of that entity, e.g., its host
range, pathogenicity, and growth conditions, likewise may be dis- OPPORTUNITY AND CHALLENGE
tinguished. If a biological species concept were adopted for all
fungi, a case could be made for multiplying the existing total num- The impression is often given that fungi in general have wider
ber of species in some groups by a factor of five or more. geographic distributions than vascular plants. In practice, the situ-
ation varies markedly between different groups. Particularly wide-
Lost Within Described Species spread are ancient fungi from phyla extending back to the first
In some groups of fungi, there has been a tradition of describing colonization of land in the Silurian region, some 300 million years
species as new if the fungus is found on a new host plant. This can before the first flowering plants. Fungi known from the early fos-
lead to the unnecessary proliferation of species names, but it also sil record include endomycorrhizas, chytrids, myxomycetes, and
can mask situations in which more than one species of a fungal some lichen-forming fungi. Also found in the category of fungi
genus occurs on a single host. When critically examined, many of with wide geographic distributions are certain nonspecific, wood-
these fungi can be keyed out based on morphological criteria rotting bracket fungi, opportunistic r-strategy spoilage molds, cop-
alone, without reference to the host. This was the case in a recent rophilous fungi, and rapidly growing soil saprobes. Detailed com-
study on Meliola in Kenya (40). The net effect of this tradition is parisons between national checklists of fungi, although incomplete,
unknown, but experience at IMI suggests that in groups such as would give some indication of the extent of overlap. However, when
Cercospora and Meliola too few, rather than too many, species are new species abound everywhere, it is difficult to know whether these
currently being recognized. Recent work on species of Alternaria discoveries are restricted, endemic species, or cosmopolitan taxa.
on Citrus exemplifies the situation in which isolates from one host The extent of our ignorance poses major problems for mycol-
are uncritically identified as a single, widely reported species con- ogists and plant pathologists working in little-studied countries or
sidered to be host specific, while, in reality, a number of distinct habitats. What is already known? Which names are correct? How
fungal species occur on that host (48). Such simplifications have does one store information on new collections? How can one be-
serious plant quarantine implications. gin to identify newly collected specimens? How does one ascer-
tain what is new? How does one gain access to available infor-
Named and Orphaned Species mation when important publications no longer in print may be
Many fungi have already been named but remain lost amongst impossible to obtain? Conversely, how does one make new infor-
the 250,000 species names that have not been characterized by mation accessible to others? These are complex problems that
modern standards. These species are not reported in the recent lit- must be addressed (23). Answers are far from easy because of the
erature and are not included in the total number of known species. lack of attention devoted to the myriad organisms involved.
Many of these obscure names are probably synonyms of known Some may say that molecular approaches are the sole answer to
species. Based on 15 fungal monographs (18), one can calculate a describing fungal biodiversity. We do not. With around 1,800
ratio of 2.5 synonyms for each “good” species. This ratio suggests fungi new to science being described every year, and not more
that of the 250,000 species names in existence 100,000 of than 100 species sequenced each year, the gap widens rather than
these are accepted species. Because only 72,000 are treated in narrows. Other than for fungi of medical and agricultural signifi-
the modern literature (21), there are 28,000 currently una- cance, the technological developments, including automated mol-
dopted “orphan” species. Although this is not a major portion of ecular and biochemical procedures, are unlikely to solve this
the absentees, it does make the total number of expected species problem within the foreseeable future. We suggest that those work-
more realistic. ing in descriptive mycology must be well supported in the essen-
When genera are studied on a world scale, comprehensive mono- tial task of characterizing newly discovered fungal species. With
graphs reveal spectacular numbers of previously undescribed spe- increased electronic resources and use of the Internet to dissemi-
cies. Of 39 species accepted in Meliolina on tropical plants, 26 nate information, the products of every mycologist describing
(67%) were new (26), and of 20 in Lichenothelia on rocks, 18 (90%) fungi can be made available rapidly along with user-friendly keys
were new (24). In some cases, the number of species known in for identification linked to diagrams explaining cryptic features and
one fungal genus is reduced significantly when species prove to illustrations of the species. Ideally, the two approaches to under-
be plurivorous. This is exemplified by an account of Didymo- standing fungal species would be combined, such that newly de-
sphaeria, in which it was determined that the 7 accepted species scribed species would be routinely sequenced and organisms rep-
had more than 100 synonyms (1). However, the usual pattern for resenting the ends of molecular cladograms would be fully described.
little-studied fungi is that when investigated a large percentage of The result would be a whole that equaled more than the sum of its
undescribed species are discovered. parts.
890 PHYTOPATHOLOGY
 The short answer to the question posed in the title, “where are dependent biotas: An overlooked aspect of conservation science. Acad.
all the undescribed fungi?”, is everywhere, including our own Sin. Bot. Bull. In press.
21. Hawksworth, D. L., Kirk, P. M., Sutton, B. C., and Pegler, D. N. 1995.
backyards. These fungi reside in unresearched niches as well as Ainsworth & Bisby’s Dictionary of the Fungi. 8th ed. CAB International,
known habitats explored by applying something other than the Wallingford, U.K.
usual isolation techniques. They exist in previously studied plants 22. Hawksworth, D. L., and Miadlikowska, J. New species of lichenicolous
and soil in developed, temperate regions, as well as the ecolog- fungi occurring on Peltigera in Ecuador and Europe. Mycol. Res. In press.
ically diverse habitats of tropical regions. Recognizing the enor- 23. Hawksworth, D. L., and Mibey, R. 1997. Information needs of inventory
mous undiscovered diversity in fungi, especially those associated programmes. Pages 55-68 in: Biodiversity Information. S. Dextre-Clarke,
P. M. Kirk, and D. L. Hawksworth, eds. CAB International, Wallingford, U.K.
with plants, soils, and insects in the tropics, is both an opportunity 24. Henssen, A. 1987. Lichenothelia, a genus of microfungi on rocks. Bibl.
and a challenge. Researchers exploring this frontier are not sur- Lichenol. 25:257-293.
prised to uncover novel fungi that often represent new orders or 25. Heywood, V. H., ed. 1995. Global Biodiversity Assessment. Cambridge
families. A vast territory ripe for exploration awaits those who ac- University Press, Cambridge.
cept the exciting challenge of discovering the world’s fungi. 26. Hughes, S. J. 1993. Meliolina and its excluded species. Mycol. Pap. 166:
1-255.
27. Hyde, K. D., Fröhlich, J., and Taylor, J. 1997. Diversity of ascomycetes
ACKNOWLEDGMENTS on palms in the tropics. Pages 141-156 in: Biodiversity of Tropical
Microfungi. K. D. Hyde, ed. Hong Kong University Press, Hong Kong.
We thank H. Gladish, M. Palm, and R. Pardo-Schultheiss for their in- 28. Kohlmeyer, J., Volkmann-Kohlmeyer, B., and Eriksson, O. E. 1996. Fun-
sightful comments and assistance in preparing this manuscript. gi on Juncus roemerianus: New marine and terrestrial ascomycetes. Mycol.
Res. 100:393-404.
LITERATURE CITED 29. Matsushima, T. 1980. Saprophytic Microfungi from Taiwan: Hyphomy-
cetes. Matsushima Mycol. Mem. 1:1-82.
1. Aptroot, A. 1995. A monograph of Didymosphaeria. Stud. Mycol. 37:1-160. 30. Matsushima, T. 1981. Saprophytic Microfungi from Taiwan. Matsushima
2. Aptroot, A., Diederich, P., Sérusiaux, E., and Sipman, H. J. M. 1997. Li- Mycol. Mem. 2:1-68.
chens and lichenicolous fungi from New Guinea. Bibl. Lichenol. 64:1- 31. Matsushima, T. 1983. Saprophytic Microfungi from Taiwan. Matsushima
220. Mycol. Mem. 3:1-90.
3. Barreto, R. W., Evans, H. C., and Ellison, C. A. 1995. The mycobiota of 32. Matsushima, T. 1985. Saprophytic Microfungi from Taiwan. Matsushima
the weed Lantana camara in Brazil, with particular reference to biologi- Mycol. Mem. 4:1-68.
cal control. Mycol. Res. 99:769-782. 33. Matsushima, T. 1987. Saprophytic Microfungi from Taiwan. Matsushima
4. Bills, G. F., and Polishook, J. D. 1994. Abundance and diversity of mi- Mycol. Mem. 5:1-100.
crofungi in leaf litter of a lowland rain forest in Costa Rica. Mycologia 34. Matsushima, T. 1989. Saprophytic Microfungi from Taiwan. Matsushima
86:187-198. Mycol. Mem. 6:1-100.
5. Brasier, C. M. 1987. The dynamics of fungal speciation. Pages 231-260 35. Matsushima, T. 1993. Saprophytic Microfungi from Taiwan. Matsushima
in: Evolutionary Biology of the Fungi. A. D. M. Rayner, C. M. Brasier, Mycol. Mem. 7:1-72, 131 pl.
and D. Moore, eds. Cambridge University Press, Cambridge. 36. Matsushima, T. 1995. Saprophytic Microfungi from Taiwan. Matsushima
6. Cannon, P. F. 1995. An ATBI—How to find one and what to do with it. Mycol. Mem. 8:1-54, 120 pl.
Inoculum 46(4):1-4. 37. Matsushima, T. 1996. Saprophytic Microfungi from Taiwan. Matsushima
7. Cannon, P. F. 1997. Diversity of the Phyllachoraceae with special ref- Mycol. Mem. 9:1-82.
erence to the tropics. Pages 255-278 in: Biodiversity of Tropical Micro- 38. Matzer, M. 1996. Lichenicolous ascomycetes with fissitunicate asci on
fungi. K. D. Hyde, ed. Hong Kong University Press, Hong Kong. foliicolous lichens. Mycol. Pap. 171:1-202.
8. Cannon, P. F., and Hawksworth, D. L. 1995. The diversity of fungi as- 39. May, R. 1994. Conceptual aspects of the quantification of the extent of
sociated with vascular plants: The known, the unknown and need to bridge biological diversity. Phil. Trans. Roy. Soc. Lond. B Biol. Sci. 345:13-20.
the knowledge gap. Adv. Plant Pathol. 11:277-302. 40. Mibey, R. K., and Hawksworth, D. L. Meliolaceae and Asterinaceae of
9. Carreiro, M. M., and Koske, R. E. 1992. Room temperature isolations the Shimba Hills, Kenya. Mycol. Pap. In press.
can bias against selection of low temperature microfungi in temperate 41. Moss, S. T., and Taylor, J. 1997. Mycobionts in the guts of millipedes—
forest soils. Mycologia 84:886-900. The Eccrinales. Mycologist 10:121-124.
10. Clay, K. 1995. Correlates of pathogen species richness in the grass fami- 42. Petrini, O., Hake, U., and Dreyfuss, M. 1990. Analysis of fungal com-
ly. Can. J. Bot. (Suppl. 1) 73:S42-S49. munities isolated from fruticose lichens. Mycologia 82:444-451.
11. Cornejo, F. H., Varela, A., and Wright, S. J. 1994. Tropical forest litter 43. Polishook, J. D., Bills, G. F., and Lodge, D. J. 1996. Microfungi from
decomposition under seasonal drought: Nutrient release, fungi and bac- decaying leaves of two rain forest trees in Puerto Rico. J. Ind. Microbiol.
teria. Oikos 70:183-190. 17:284-294.
12. Da Silva, M., and Minter, D. W. 1995. Fungi from Brazil recorded by 44. Rambelli, A., Persiani, A. M., Maggi, O., Lunghini, D., Onofri, S., Riess,
Batista and co-workers. Mycol. Pap. 169:1-585. S., Dowgiallo, G., and Puppi, G. 1983. Comparative studies on micro-
13. Diederich, P. 1995. The lichenicolous heterobasidiomycetes. Bibl. Lichenol. fungi in tropical ecosystems. Mycological Studies in South Western Ivory
61:1-198. Coast Forest. UNESCO, Rome.
14. Domsch, K. H., Gams, W., and Anderson, T.-H. 1980. Compendium of 45. Rossman, A. Y. 1995. A strategy for an all-taxa inventory of fungal
Soil Fungi. Vol. 1. Academic Press, New York. biodiversity. Pages 169-194 in: Biodiversity and Terrestrial Ecosystems.
15. Farr, D. F., Bills, G. F., Chamuris, G. P., and Rossman, A. Y. 1989. Fungi C. Peng and C. H. Chou, eds. Academia Sinica, Taiwan.
on Plants and Plant Products in the United States. The American Phyto- 46. Sankaran, K. V., Sutton, B. C., and Minter, D. W. 1995. A checklist of
pathological Society, St. Paul, MN. fungi recorded on Eucalyptus. Mycol. Pap. 170:1-376.
16. Hammond, P. M. 1992. Species inventory. Pages 17-39 in: Global Biodi- 47. Shivas, R. G., and Hyde, K. D. 1997. Biodiversity of plant pathogenic
versity. B. Groombridge, ed. Chapman and Hall, London. fungi in the tropics. Pages 47-56 in: Biodiversity of Tropical Microfungi.
17. Hawksworth, D. L. 1991. The fungal dimension of biodiversity: Magni- K. D. Hyde, ed. Hong Kong University Press, Hong Kong.
tude, significance, and conservation. Mycol. Res. 95:641-655. 48. Simmons, E. 1990. Alternaria themes and variations (27-53). Myco-
18. Hawksworth, D. L. 1992. The need for a more effective biological no- taxon 37:79-119.
menclature for the 21st century. Bot. J. Linn. Soc. 109:543-567. 49. Thorn, R. G., Reddy, C. A., Harris, D., and Paul, E. A. 1996. Isolation of
19. Hawksworth, D. L. 1993. The tropical fungal biota: Census, pertinence, saprophytic basidiomycetes from soil. Appl. Environ. Microbiol. 62:4288-
prophylaxis, and prognosis. Pages 265-293 in: Aspects of Tropical My- 4292.
cology. S. Isaac, J. S. Frankland, R. Watling, and A. J. S. Whalley, eds. 50. Weir, A., and Hammond, P. M. 1997. Laboulbeniales on beetles: Host
Cambridge University Press, Cambridge. utilisation patterns and species richness of the parasites. Biodivers. Conserv.
20. Hawksworth, D. L. The consequences of plant extinctions for their 6:701-719.

Vol. 87, No. 9, 1997 891