Mycol. Res.

9S (6): 641--655 (1991) Printed in Great Britain 641

Presidential address 1990
The fungal dimension of biodiversity: magnitude, significance,
and conservation
D. L. HAWKSWORTH
International Mycological Institute, Kew, Surrey TW9 3AF, UK
Fungi, members of the kingdoms Chromista, Fungi S.str. and Protozoa studied by mycologists, have received scant consideration in
discussions on biodiversity. The number of known species is about 69000, but that in the world is conservatively estimated at
1'5 million; six-times higher than hitherto suggested. The new world estimate is primarily based on vascular plant:fungus ratios in
different regions. It is considered conservative as: (1) it is based on the lower estimates of world vascular plants; (2) no separate
provision is made for the vast numbers of insects now suggested to exist; (3) ratios are based on areas still not fully known
mycologically; and (4) no allowance is made for higher ratios in tropical and polar regions. Evidence that numerous new species
remain to be found is presented. This realization has major implications for systematic manpower, resources, and classification. Fungi
have and continue to playa vital role in the evolution of terrestrial life (especially through mutualisms), ecosystem function and the
maintenance of biodiversity, human progress, and the operation of Gaia. Conservation in situ and ex situ are complementary, and the
significance of culture collections is stressed. International collaboration is required to develop a world inventory, quantify functional
roles, and for effective conservation.
'Biodiversity', the extent of biological variation on Earth, has
come to the fore as a key issue in science and politics for the
1990s. First used as 'BioDiversity' in the title of a scientific
meeting in Washington, D.C. in 1986 (Wilson, 1988: p. v), it
has been rapidly adopted as a contraction of 'biotic diversity'
and 'biological diversity'. Interest has been inflamed by
concern over the conservation of genetic resources, destrudion
of forests, extinction of species, and the effects of global
warming. A plethora of texts and reports has resulted; some
of the more significant since 1985 are Norton (1986a), Soule
(1986), U.S. Congress Office of Technology Assessment
(1987), Wolf (1987), Cronk (1988), Lugo (1988), Wilson
(1988), Knutson & Stoner (1989), U.s. National Science Board
(1989), di Castri & Younes (1990), Keystone Center (1990),
McNeely et al. (1990), and u.s. Board on Agriculture
(1991).
While many of the principles and discussions of broader
issues raised in these works are relevant to mycology, most
lack any substantive content on fungi, or indeed in many cases
on any micro-organisms. Exceptions with sections on at least
some micro-organism aspects are: U.s. Congress Office of
Technology Assessment (1987), Knutson & Stoner (1989),
U.S. National Science Board (1989), di Castri & Younes (1990),
and U.S. Board on Agriculture (1991).
The aim of this address is to broaden the biodiversity
debate by focusing on its fungal dimension; the magnitude of
the task and its implications for systematics; the significance
of fungi in evolution, ecosystem function, human progress,
and to Gaia; and the conservation of fungi. Biodiversity can
be explored at a variety of levels: in terms of ecosystems,
41
species, or populations. Knowledge of all of these is pertinent
to a thorough appreciation of the fungal dimension, but here
I will centre on species biodiversity; that is basal to discussions
at other levels.
DAVID L. HAWKSWORTH
President, British Mycological Society, 1990
MYC 95
Presidential address
642
MAGNITUDE
Circumscription
What are fungi? An answer is fundamental to a consideration
of their magnitude. In the first half of this century, 'Fungi'
were generally treated as a part of the kingdom 'Plantae' in
the subdivision 'Thallophyta' and placed alongside Bacteria,
Lichenes, and Algae (e.g. Fitzpatrick 1930: p. 2). The five-
kingdom system of Whittaker (1969) was rapidly adopted
which accepted a kingdom 'Fungi' (including myxomycetes
and oomycetes) as distinct from Animalia, Monera (bacteria
and cyanobacteria), Plantae (including bryophytes and non-
flagellate algae), and Prostista [i.e. 'Protoctista 'J (including
plasmodiophoromycetes and hyphochytridiomycetes). It is
now clear that this break from tradition did not go far enough
in representing the diversity of life. Greater numbers of
kingdoms and arrangements within those have been proposed
by a variety of workers in the 1980s (e.g. Cavalier-Smith,
1981, 1987; Corliss, 1984; Tehler, 1988; von Arx, 1987).
At present, the prevailing view, based on a combination of
the limited ultrastructural and molecular data now available,
is to distribute the phyla (i.e. 'divisions ') generally studied
by mycologists between the kingdoms Protozoa (incl.
Myxomycota), Chromista (incl. Oomycota), and Fungi 5.5tr.
An attempt to reflect the current view of the overall
relationships and phylogeny of the phyla concerned is
incorporated into Fig. 1. However. knowledge in this area is
advancing rapidly, and it has even been suggested that the
whole 'kingdom' concept in eukaryotes needs rethinking
(Sogin et aI., 1989).
In the remainder of this address, 'fungi' is used in the
admittedly arbitrary but traditional sense of 'organisms
studied by mycologists', i.e. to encompass those now also
placed in other kingdoms. The inclusion of lichen-forming
fungi in the sixth edition of the Dictionary of the Fungi
(Ainsworth et al., 1971), and in the Index of Fungi from 1970,
at first shocked some. However, a thorough integration was
the only logical possibility, especially in view of the diversity
of biologies involved (Hawksworth, 1988a); this is now
accepted as the norm and does not merit further comment
here. Unless otherwise restricted, 'fungi' as used in this
address encompasses those forming lichens.
Described species
The best estimates for the number of known species of fungi
come from the additions of totals of accepted species given
for each genus in the first and seventh editions of the
Dictionary. These were the only two where counting was
undertaken from individual entries and gave about 38000
(Ainsworth & Bisby, 1943: p. 204; Bisby & Ainsworth, 1943)
and 64200 species (Hawksworth et al., 1983: p.266)
respectively.
On average 700 species were described as new to science
each year from 1920 to 1950 (Ainsworth, 1954). The annual
total catalogued in the Index of Fungi reached around 1400 in
1961 (Ainsworth, 1963), 1500 by 1968 (Ainsworth, 1968),
and has averaged 1700 each year for 1986-90 (p. M. Kirk, pers.
comm.); that the pre-1970 issues did not cover Iichenized
species only partly accounts for this ' rise' (see below).
Taking the 1700 annual rate, it follows that approximately
BASIDIOMYCOTA
_________ Fruil body retained
The PHYLA of "FUNGI"
- - - - - - - - - Meiotic exospores fonned (basidiosporcs)
--------- Fruit body formed
USTOMYCOTA
UREDINIOMYCOTA
_________ Dikaryon formed
Meiotic endospores fanned (ascnspores)
Zygospore lost
_________ Mitospores produced
Cilia lost
Zygospore developed
- - - - - - - - - Chi.i" cell wall
Loss of amoeboid trophic phase
- - - - - - - - - Cilia developed
_________ Chromosomc!i formed
Mitochondria originate symhiotically
-
"
,
,
,
,
,
,
,
,
,
,
,
,
- , -
, --
..... ~ ,
,
,
Fig. 1. Schematic representation of selected characteristics and probable relationships of the phyla of 'fungi', and the kingdoms to
which they are now assigned. The Eubacteria are included only for reference.
D. L. Hawksworth 643
Revised estimates. Four decades after Martin's (195 I) estimate,
and to improve the mycological input to the biodiversity
1 Hawksworth et aI. (1983).
2 Vol. 5 (6-20); twice-yearly, International Mycological Institute.
3 2'5: 1 synonymy assumed.
II 900 'new' species have been described since 1983.
Assuming that as in the past 'more than half' (Ainsworth &
Ciferri, 1955) or 'one out of three' (Bisby & Ainsworth, 1943)
can be expected to be unnecessary synonyms, it seems not
unreasonable to accept a synonymy level of 2'5: 1. On that
basis, the number of described species accepted in the latest
Dictionary should be increased by 4800 to up-date it to the
end of 1990. Making no allowance for double counting due to
separately named anamorphs of known teleomorphs (see
below), this provides a revised figure of 69000 for the number
of species of fungi currently known (Table I).
Undescribed species
Previous estimates. Fries (1825: p. 47) considered that the
fungi would prove to be the largest group in the 'orbis
vegetabilia', analogous to the insects in the' animalia'; that it
would be easy to make mycology larger than the rest of
botany (Fries, 1828: p. 107); and that describing them
individually would frustrate the memory of man (Fries, 1829:
p. vii). While, as far as I have been able to discover, he
published no overall numerical estimate, he did indicate that
the number of agaric species could well be as much as 40000
(Fries, 1849: pp. 267-268), the pyrenomycetes alone 100000
(lac. cit. p. 378), and that science could collapse if all foliicolous
species were described (/oc. cit. p. 509). On the basis of these
statements, I suspect he would have considered an estimate of
250000 low. According to Cooke (1895: p.319), in 1872
A. de Bary went for a more modest ISO000 species.
Bisby & Ainsworth (1943: p. 18) made a 'guess' that about
one third of the species were then known, Le. 'that there are
about 100000 species'. Based on an analysis of fungal data in
Iowa in relation to the numbers of vascular plants, Martin
(1951: p. 177) regarded the figure of 100000 'excessively
conservative and that the total number may be of substantially
the same order of magnitude as the number of species of
vascular plants', then estimated at about 260000 Gones,
1951). Ainsworth (1968: p.513) considered this 'probably
still on the conservative side', noting that the regularity in the
pattern of description of new species found was as to be
expected if those 'proposed each year are a random sample
from a large undescribed population '.
This issue has not attracted much subsequent attention,
although Korf (1991) considered that at least half and
probably more than two thirds of the world's discomycetes
remained undescribed.
A:B
Vascular Plants
1
Fungi
2
(A) (B) Region
debate, it is opportune to calculate an updated figure.
Intensive 'knock-down' studies of the insects associated with
individual trees in the tropics have provided the basis for
recalculations of the extent of insect diversity. Extrapolations
have led to estimates of the world's insect species as in the
range 10-80000000 (Stork, 1988). While each step in such
arguments can be challenged, especially the extent of
insect-host specificity in the tropics (May, 1988), the number
of insects is likely to be at least 6000000 (R. M. May,
unpub!.; Thomas, 1990).
In order to provide new estimates in which we can have
confidence, it is vital to base these on the best information
available. In the case of the fungi, the most intensively studied
region in the world is the British Isles. Bisby & Ainsworth
(1943) considered that there were perhaps then 6000 'good'
fungus species recognized; however, they did not allow for
lichen-forming species which at that time would have added
1400 further species (Watson, 1953). The total figure in
1943/53 would then have been 7400. Today that sum stands
at around 12 000 (Sims et al., 1988), an increase of 62 % in
47 years, and a rate of 13 % per decade. The total of accepted
British ascomycdes alone is currently 5100 species (Cannon
et al., 1985), and a staggering 4931 fungi are known from
y orkshire (Table 2).
The ratio between the number of vascular plants and fungi
from all substrata (not only plants and plant products) in the
British Isles as a whole is now about I: 6; i.e. about twice that
in 1943/53. In the better-studied counties, islands, or sites it
is around I: 3 or I: 4 (Table 2). This difference is to be
expected as no site in Britain can yet claim to be fully recorded
for fungi. Indeed the high Yorkshire figure is certainly due to
Table 2. Comparison of the numbers of vascular plants and fungi
occurring in well-studied sites in the British Isles
British Isles 2089
3
12000' 1: 6
Warwickshire 1231' 2795
6
1:2
Yorkshire 1314' 4931
6
1:4
Hebrides 860 3769'0 1: 4
Isle of Mull 783" 2760
12
1: 4
Slapton 490
13
1619
14
1:3
Wheatfen Broad 238
15
996
15
1: 4
1 Pteridophytes, gymnosperms and spermatophytes, not garden species
unless common escapes; excluding 'microspecies' and hybrids.
2 In all habitats, including lichen-forming species, and not double-counting
anamorphs with known teleomorphs (except for 'British Isles).
3 Clapham et al. (1987); S. L. Jury (pers. comm.); excluding ca. 25000
species in gardens U. C. Alexander & S. L. Jury, pers. comm.).
Sims et aI. (1968).
, Cadbury el aI. (1971).
6 Clark (1980, 1986).
, M. R. D. Seaward (pers. comm.).
8 Bramley (1985); Seaward (1989, in lill.).
9 M.RD. Seaward (pers. comm.).
10 Dennis (1986); M. R. D. Seaward (pers. comm.).
11 Bangerter & Cannon (1978); Bangerter el aI. (1978); Clark & Jermy
(1990).
12 James (1976); Watling (1978); Henderson & Watling (1985).
13 Brookes & Burns (1969).
" Hawksworth (1986, unpub!.).
15 M. B. Ellis (in lill.).
Species
64200
4600
3
69000
Dictionary of the Fungi 1983
'
Index of Fungi 1983-90
2
Total
Source
Table 1. The number of known species of fungi
41-2
Presidential address 644
Table 3. Estimates of the total number of species of fungi in the world
derived by different methods (see text for further explanation)
alone on 34 vascular plant species in an alpine sedge
community; a ratio of 1: 3'8. That ratio could not unreasonably
be expected to be as much as 1: 6 had all fungi (including rusts,
other ascomycetes and conidial fungi) been taken into account.
Extrapolated to the world that would give 1620000 [Estimate
0] on vascular plants alone.
Based on less detailed enquiries, I previously and hesitantly
suggested that there might be 800000 species of fungi in the
world (in di Castri & Younes, 1990). The data now presented
demonstrate that figure to be an underestimate. A mean of
Estimates A-O (admittedly not comparable) yields a figure of
1262250 [Estimate E; Table 3]; as Estimates C and 0 are not
based on all substrata, B is certainly based on inadequate
study, and hosts and additional fungi continue to be found in
all these areas, it may seem not unreasonable to suggest
1650000 [Estimate F] as a conservative figure for the actual
number of species of fungi in the world.
However, all the preceding calculations do not allow for
anamorphs where the corresponding teleomorphs are named,
and so are also included. Bisby & Ainsworth (1943) made a
reduction of one third in the anamorph species totals to allow
for this factor. If the proportions between the different groups
revealed by the seventh edition of the Dictionary are assumed
still to hold when all fungi are known, conidial fungi would
represent about 26'5 % of the total. Applied to Estimate F, that
proVides a figure of 437750 species of conidial fungi; a one
third reduction therefore necessitates a cut of 145750 species.
In the case of Estimate F, that proVides a corrected figure of
1504800 species [Estimate G]. This is the number which I
suggest, rounded to 1'5 million, is used as the best estimate of
the number of species of fungi in the world.
I consider that 1'5 million, although six times greater than
any previously proposed by other authors, will nevertheless
prove to be conservative for four reasons:
(i) A world total of 270000 species of vascular plants has
been assumed, even though estimates up to 400000 have
been published (Wolf, 1987). If the higher estimate were
correct, that would provide a figure of 2400000 fungi.
(ii) No separate allowance has been made for fungi that
might occur on the vast numbers of insects now postulated;
that ecological niche has scarcely been studied in the British
Isles and is therefore not adequately represented in the data
set. Of museum specimens of 1100 insect species from eastern
Fennoscandia, 166 (15 %) supported 88 members of the
that county having been studied by more mycologists than
any other over the years, 65 as opposed to 35 in the next
highest, Surrey (G. C. Ainsworth, in lift.). This interpretation is
also consistent with Dennis (1986: p. 9) who after 35 years
examining Hebridean fungi considered' ... it would be unwise
to imagine the list represents more than perhaps half of the
species actually present '. For example, many species in a site
on mosses, lichens, in soil, on insects, in water, etc, which are
known elsewhere in the British Isles will be present but remain
undetected unless the appropriate techniques and specialists
are employed.
Although the ratio presented for the British Isles does not
take into account 'garden species', most of the cultivated
species and casual aliens, of vascular plants, but does
incorporate fungi from them, I believe the 1: 6 ratio can be
sustained as: (a) the British fungus flora is still underworked,
as evidenced by the continuing flow of new records (see
above); and (b) cultivated' garden species' generally have few
of the specific fungi which occur on them in regions where
they are native, indeed quarantine procedures aim to eliminate
them. If a ratio of 1: 6 is accepted and extrapolated to the
world, assuming a conservative 270000 vascular plants yields
an estimate of 1620000 species of fungi [Estimate A].
Finland, which has received considerable but much less
mycological attention, has 1350 vascular plants and about
5900 fungi (Rassi & Viisiiinen, 1987; T. Ahti. pers. comm.),
giving a ratio of 1: 4. Similarly in Switzerland there are about
2500 vascular plants and 10000 fungi recorded (E. Miiller, in
lift.) yielding a ratio of 1: 4. These data add weight to the view
that the British data are not atypical, at least for Europe.
In the u.s.A. (including Alaska, Hawaii, Puerto Rico, and
the Virgin Islands), 8792 vascular plants have 12666 fungi on
them and their products (Farr et al., 1989). While this gives a
ratio of 1: 1'4, as 21500 plants are actually known in the
region 0. Kartesz, in lift.) that should be revised down to
1: 0'6. However, the fungi on native vascular plants, especially
microfungi on species in the tropical U.s.A., have scarcely
been investigated. How many fungi have been found on other
substrata and hosts in the country has not been catalogued,
but can be expected to include about 4000 lichens (3409 occur
in the continental u.s.A. and Canada alone; Egan, 1987), and
4000 from the remainder. That gives an approximate recorded
total of about 20700 species and a ratio of only about 1: 1. A
world extrapolation based on 270000 vascular plants then
gives only 270000 [Estimate B].
No comparable total for fungi on plants and plant products
is available for the British Isles. However, the authors of 134
of the accounts of individual plants in the series 'Biological
Flora of the British Isles' Uournal of Ecology 29-77, 1948-89)
have, to varying extents, attempted to catalogue associated
fungi; these data were mainly taken from literature sources
and not based on critical mycological studies. The number per
host listed varied from 0 to 81, with a mean of 8'5. Not all
were restricted to a single host and if one third are taken not
to be host specific, a mean of 5'7 results. Extrapolated to the
world that would give 1539000 fungi on vascular plants alone
[Estimate C]. Some indication that this data set is not
exceptional comes from the study of Nograsek (1990) who
found 128 ascomycetes (excluding inoperculate discomycetes)
Estimate Basis
A British Isles
B US 'Plants and Plant Products'
C 'Biological Flora of the British Isles'
D Alpine sedge community
E Mean A-D
F Plus allowance for unstudied substrata
G Minus allowance for anamorphs having
known teleomorphs
H Assuming 30 million insects
Ratio
1:6
1:1
1:5'7
1:6
Total
species
nwnber
1620000
270000
1539000
1620000
1262250
1650000
1504800
3004800
D. L. Hawksworth
Laboulbeniales (Hulden, 1983); that figure must be an
underestimate of those in nature because of the sampling
method. As there are about 9850 species of Coleoptera and
Diptera in the British Isles (Sims et aI., 1988), it might not be
unreasonable to expect 750 species in that order alone rather
than the 45 recorded (Cannon et al., 1985). If only 5 % of insect
species had obligate mutualistic symbionts, parasites, or
commensals, that would give estimates of: 250000 fungi on
5 million insects, 500000 fungi on 10 million insects, and
1500000 on 30 million. Were the last figure correct, the
fungal total could be as high as 3004800 species [Estimate H].
If a 15 % rate were assumed on the highest insect estimate, a
highly speculative 13'5 million fungi emerges.
(iii) The ratios of vascular plants: fungi used in deriving the
estimates can be expected to be too low when all possible
fungal habitats have been exhaustively studied. In the case of
the British Isles, the total number of fungi recorded is rising at
the rate of about 13 %each decade (see above), and there is no
sign of it diminishing, while that for vascular plants remains
almost constant; on that basis by the year 2000 the ratio
would be almost 1: 7 (13 600 fungi), by 2010 1: 7'5 (15100
fungi), etc. Any critical study of an area in the British Isles
yields new records; about 60 fungi have been added to the
national list during the Slapton survey alone.
(iv) Ratios derived almost entirely from North Temperate
regions have been assumed to apply world-wide, but could be
higher both in the tropics and polar regions. Although no
country from either of these areas has been studied sufficiently
intensively to proVide information comparable to that for the
British Isles, there are indications why higher ratios might be
expected in both.
In India, there are 15000 vascular plants (Nayar, 1989) and
either 15500 (Singh, 1980; Sarbhoy et al., 1986) or 23000
(Nayar, 1989) fungi yielding ratios of 1: 1 or 1: 1'5. However,
Subramanian (1986) provided evidence of the substantial
numbers of new species still to be found in India; for example
10 (43 %) of 23 coronophoraceous fungi from the Western
Ghats alone proved to be undescribed. In Sierra Leone, data
supplied by F. C. Deighton (in litt.) provide a ratio of 1:0'5 if
all vascular plants are totalled (about 4100 species), but 1:2 if
only those collected from are considered. In Papua New
Guinea, there are 2390 species of fungi treated in Shaw (1984)
and about 13000 vascular plants (Womersley, 1978; G.
Guymer, in litt.), giving an even lower ratio of 1 :0'2. None of
these areas has been sufficiently well-studied to enable any
firm conclusions to be drawn.
It has been estimated that two thirds of the world's
flowering plants occur in the tropics (Raven, 1988), and the
proportion for insects could be much more in that region if the
results of studies such as that of Stork (1988) are upheld. In the
absence of non-speculative data it would be unsound to raise
the vascular plant: fungus ratio on that basis. While it might
reasonably be expected that fungi on perennial leaves (with
associated fungicolous species) and the numbers of endophytic
and entomogenous fungi could significantly affect that ratio,
at present it cannot be confidently asserted that vascular plant
host specificity occurs to the same extent in tropical as in
temperate forests. It is also pOSSible that the lichen-forming
species, at least on rocks and the ground, are proportionately
645
fewer in the tropics. In order to obtain sounder data,
mycologists (and other microbiologists) should fonn an
integral part of tropical survey teams, as recommended
by the U.S. Strategy Conference on Biological Diversity
(1982: p. 91). In-depth site studies are crucial to placing
estimates on a finner base (U.S. National Science Board, 1989:
p.9).
In boreal to arctic and antarctic habitats where vascular
plant numbers are low, the ratio will also be much higher. In
Antarctica and the subantarctic islands where lichen-fonning
fungi are especially important components of the vegetation,
72 vascular plants compares with at least 529 fungi (Dodge,
1974; Pegler et al., 1980; Pugh & Allsopp, 1982) giving 1: 7;
excluding the antarctic islands, with only two vascular plants
that ratio on current data appears to be about 1: 100, even
though the non-lichenized fungi have scarcely been in-
vestigated. 'Hot' deserts may follow a similar pattern to cold
ones, especially where lichen crusts are a major component of
the ground cover, and if soil fungi are selectively isolated.
However, some fungi, for example lichen-fonning and
ruderal species, appear to have rather wide geographical
distributions as compared to vascular plants. If these were a
majority, this would be expected to reduce all calculated ratios
based on described species with increases in the size of the
geographic region considered. In view of the scant data on
fungal biogeography and endemism, a meaningful quantitative
allowance cannot be made for this tendency at present. In
view of discussions of the first three factors above, it
nevertheless seems not unreasonable to assume that they will
more than compensate for any variances in the 1'5 million due
to any overall 'geographical' factor.
Additional evidence. If Estimate G of 1'5 million species
approximates to the real situation, only 4'6% of the world's
fungi have so far been recognized. If that hypothesis is correct,
vast numbers of fungi must remain to be discovered. Evidence
that this is the case is provided by a variety of observations
and data sets, examples of which are presented here.
Tropical regions would be expected to be the richest
sources of new species. In Brazil, A. C. Batista (1916-67) and
his co-workers described approximately 3500 species between
1954 and 1972, mostly in the series Publicafoes Instituto de
Micologia Universidade do Recife. These new fungi came mainly
from easily accessible parts of the Amazon, and several species
new to science were regularly discovered from single perennial
leaves. Singer (1989) published 276 new species of agarics of
which 241 were from Central and South America. Particularly
tellingly, Rogerson et al. (1990) found 10 of 242 species of
fungi collected by a non-specialist in the Guyana Highlands to
be new.
This pattern is reflected in other continents. In Sierra Leone,
of 1848 species mostly collected by F. C. Deighton in
1926-55, 554 (30%) have been described as new and many
more in IMI await fonnal description (F. C. Deighton, in lift.).
In East Africa, of 628 macrolichens recognized by Swinscow
& Krog (1988), 79 species (13 %) had been described as new
from their material collected in 1969-77; and of 389 agarics
accepted by Pegler (1977) 63 species (16%) were new. In the
Lesser Antilles, 70 (15 %) of the 457 species of agarics were
Presidential address
646
Table 4. Decadal rate of increase in numbers of described species in
selected habitats
new species of ascomycetes were found on 52 specimens of
the Australasian and south-east Asian moss genus Dawsonia
that chanced to be in the herbarium at Munchen (Dobbeler,
1981); and the symbionts of hepatics are scarcely studied
(Boullard, 1988). Fungi growing on fungi (fungicolous fungi),
especially in the tropics, receive equally poor attention: 774
conidial fungi alone occur on other fungi (Hawksworth, 1981);
19 (32 %) of 60 fungicolous (including lichenicolous) nec-
triaceous species, mainly from the tropics, studied by Samuels
(1988) were new; and 120 new species have been described
from other fungi in Sierra Leone (F. C. Deighton, in litt.). The
same pattern is to be expected for fungi on algae (algicolous
fungi), especially in non-marine habitats, and this appears to
be true at least for chytridiaceous species on tropical heshwater
algae (G. A. Beakes, pers. comm.).
Fungi on and in insects and other arthropods are remarkably
little studied, and it is difficult to be certain how host specific
many are. The Laboulbeniales are perhaps the largest group,
occurring mainly on flies and beetles; Hulden (1983) found
that 24 (27%) of 88 species in eastern Fennoscandia were new.
Each species of thrips appears to have a unique Hirsutella (H. C.
Evans, pers. camm.), and Cordyceps species also evidently
have narrow host ranges to judge hom the 224 species now
known (Kobayasi & Shimizu, 1983). An indication of the
richness of entomogenous fungi in the tropics is Petch's (1921)
discovery of 60 Hypocrella species on scale insects just in Sri
Lanka. Yeast-like mutualistic fungi in the guts of wood-boring
beetles and also Trichomycetes in the digestive tracts of
insects are hardly known; when adequately studied numerous
new entomogenous species can be expected. Lichtwardt
(1986: p. 42) noted that while trichomycete species appeared
to often be able to infest more than one insect species or
genus, they could be restricted at the family level.
Nematophagous fungi, both endoparasitic and trapping,
also remain scarcely studied outside temperate regions (Kerry,
1984) and can be expected to be another rich source of novel
fungi.
The fungi on vascular plants, especially native rather than
crop species, are inadequately known even in Europe.
Nograsek (1990) found 28 (22 %) of 128 ascomycete species
in a sedge community in the Alps to be new to science (see
also above). It is still far from clear how many fungi can be
expected to be restricted to particular host vascular plants.
The numbers certainly vary widely between hosts, and in the
case of crop plants while large numbers may be known (e.g.
about 200 known only on sugarcane; Sivanesan & Waller,
1986), how many also may occur on native plants is uncertain.
Hosts with no recorded fungi Widespread as weeds or in
gardens when studied in their native habitats repeatedly yield
novel species; for example, Evans et al. (l991) discovered four
undescribed (Pegler, 1983). In Papua New Guinea, Shaw
(1984) reported that 30 genera and 604 (25 %) of 2390 species
of fungi had been based on material from the country, many
collected in 1963-77. Growth in our knowledge of Northern
Hemisphere fungi also remains grossly inadequate; for
example, 25 (15 %) of 168 hyphomycetes collected in Manitoba
and Saskatchewan in 1965-69 were found to be new (Sutton,
1973), and 61 new fungi to science were found during the
Slapton and Warwickshire studies in the British Isles (Clark,
1980; Hawksworth, 1986).
Critical world or even regional monographic studies can
yield remarkably high numbers of new species; 95 (85 %) of
112 veiled species of Hebeloma in the western u.s.A. (Smith
et aI., 1983); 30 (86%) of 35 species of Oropogon mainly from
the New World (Esslinger, 1989); 100 (71 %) of 140 bolete
species in Malaysia (Comer, 1972); 24 (29%) of 84 species of
Meliola in the State of Kerala in India (Hosagoudar & Goos,
1990); 29 (57%) of 51 hygrophoraceous fungi in New
Zealand (Horak, 1990); etc. Even traditionally well-studied
genera are not exempt hom this trend; 123 new species names
in Penicillium have been introduced since the monograph
of Pitt (1980), approaching a doubling of the 150 then
accepted.
Little-explored habitats are a major source of novel fungi
world-wide. The number of aquatic hyphomycetes, 'Ingoldian
fungi', now stands at 261 species O. Webster, in litt.), all but
38 of which (i.e. 85 %) have been described since 1960. That
of marine fungi has risen hom 209 species in 1979 to 321 in
1990 (Kohlmeyer & Volkmann-Kohlmeyer, 1991). That of
foliicolous lichens from 236 in 1952 to 345 in 1985, all but
two of the additional 109 species being recognized since 1970
(Farkas, 1986).
The fungi obligately growing on lichens (lichenicolous
fungi) have proved to be remarkably rich in novel taxa.
The number known increased from 457 species in 1976 to 682
in 1989 (Clauzade et al., 1989); that number continues to
escallate. Of 124 species reported from Greenland, 24 (19%)
were new to science (Alstrup & Hawksworth, 1990); 14 (24%)
of 58 species on epiphytic lichens in Luxembourg were
undescribed (Diederich, 1989); 10 (22 %) of 45 species on
lecideioid lichens were also new (Triebel, 1989). Some lichen
. hosts are remarkably rich in these special fungi; 82 species
occur on Peltigera, of which 52 are known only on that genus
(Hawksworth, 1980; Alstrup, in litt.). I now consider my
earlier estimate of 1000 lichenicolous species (Hawksworth,
1982 a) to be too low, and the number of obligately
lichenicolous fungi to be at least 2000 (compared with around
13 500 lichenized species); that total substantially exceeds
those of the known Oomycota, Zygomycota, gastroid fungi,
Ustomycota, or Myxomycota.
There are indications that the above pattern of explosive
increases in the number of species known in such niches, at a
rate of 20-49% per decade (Table 4), would be seen in others
were sufficient numbers of workers to tum their attention to
them. For example, fungi on bryophytes (mosses and
liverworts): Dobbeler (1978) described 9 (27%) new genera
out of 33 accepted, and 62 (50%) new species amongst 123
pyrenomycetes recognized on their gametophytes, most
material coming from Central Europe; two new genera and 21
Habitat
Aquatic hyphomycetes
Foliicolous lichens
Lichenicolous fungi
Marine fungi
Decadal increase
(%)
28
20
38
49
D. L. Hawksworth
new species (two also new genera) on the 2-3 m tall Mimosa
pigra in Mexico. That many hosts with reported species
remain most inadequately studied can be illustrated by
numerous examples: Holm & Holm (1981) found six new
ascomycetes amongst 24 on five Lycopodium species in the
Nordic countries; B. C. Sutton and his co-workers have
described 18 new genera and 55 new species of conidial fungi
alone on Eucalyptus litter since 1973, and have at least 16 more
to publish (B. C. Sutton, pers. comm.); this pattern is repeated
in studies on Castanea fallen cupules, Juniperus, fallen Laurus
leaves, etc.
Dung is also most inadequately studied. Lundqvist (1972)
found that of 100 species of the primarily coprophilous
Sordariaceae s. lat. in the Nordic countries 28 (28%) were new
to science. While soil isolations have been a focus of attention
for screening by commercial concerns, the fungi obtained are
rarely critically identified. Soils also, however, are a further
source of undescribed species and even in genera such as
Penicillium apparently local species are still being discovered;
25 described since 1980 from soil are currently accepted
(Z. Kozakiewicz, pers. comm.).
Further, previously unsuspected habitats with novel fungi
continue to come to light. For example, the' cryptoendolithic'
communities immersed in rock in Antarctica (Friedmann,
1982), and the anaerobic flagellate fungi in the guts of
ruminant mammals (Orpin, 1988). Potentially the largest
untapped pool is almost certainly endophytic fungi forming
mutualistic associations with the aerial parts of vascular
plants; 'mycophyllas' (Lewis, 1987). It has been suggested
that mycophyllas may be as frequent as mycorrhizas (Carroll,
1988), and even that each tropical tree species has 3-4
characteristic endophytic fungi (M. M. Dreyfuss, pers. comm.).
How many of such fungi are new rather than morphs of ones
fruiting on the surfaces, or after the hosts die on their debris,
remains uncertain.
Perhaps most instructive overall is that the numbers of new
species catalogued in the Index of Fungi are far from declining
and that the rate has increased (see above). The volume
(excluding cumulative indexes) covering the years 1981-90 is
1103 pages in length, compared with 649 pages for 1971-80;
lichen-forming fungi were covered in both these decades. The
key factor limiting the number of new species described is the
number of systematic mycologists available to undertake
such work. At the International Mycological Institute, for
example, I believe that we have material of about 1000 species
awaiting description; a pattern reflected in mycological
laboratories and herbaria throughout the world.
Implications for systematics
Manpower. The appreciation that only 4'6% of the world's
fungi are currently known has major implications for
systematic mycology. It has taken the 261 years since Micheli
(1729) made his attempt at a critical compilation of the known
fungi to reach this point. At the current rate of 1700 species
per year and even assuming all were 'good' (see above),
the remaining 1435 800 species would take 844 years. The
scale of the research effort to significantly reduce that time
scale would be enormous. To complete the inventory in 100
647
years would involve increasing the existing systematic
manpower by over eight times, and to achieve this by the year
2000 a staggering 80 times.
The demand for the products of systematics is currently
greater than it has ever been (Hawksworth & Bisby, 1988).
This arises from a renewed interest in conservation, bio-
diversity, the environment. and biotechnology at the
profeSSional scientist, amateur, and public levels. The growth
in membership of charities and public organizations in the
conservation arena since 1970 has been remarkable (Hodge,
1990). This situation has arisen at a time when the number of
taxonomic positions in developed countries is in decline.
Burdsall (1990) notes that in the U.S.A. the number of
systematic mycological positions in three key institutions has
declined from 13 to 6 in recent years. In the UK, while this
situation has not been mirrored in the prinCiple mycological
institutions, there is now no university department in the
country with a head or personal chair in systematic mycology.
This has grave implications for mycological training, sys-
tematic aspects of which are increasingly falling to other
institutions, including IML
Such institutions have, however, progressively come under
financial pressure to focus on revenue-generating activities.
While these are not necessarily incompatible with sound
systematic work and indeed can enable resources to be
chanelled to areas of greatest need, the time many professional
taxonomists can devote to the description and illustration of
species which are not yet known to be of importance is
curtailed.
It is imperative that the time those with taxonomic skills
can employ in this field is utilized to maximum effect.
Particularly disturbing is the time it often takes to unravel
legalistic nomenclatural problems, or to obtain original material
of inadequately described long-forgotten species names that
just might be the one in hand. To describe a species as new is
much less time consuming than finding if it has already been
named, but perhaps in an incorrect genus, family, or group.
There is a need for more taxonomy and less nomenclature;
indeed this complements improved nomenclatural stability as
one of the two main driving forces behind the current
international move to establish Lists of Names in Current Use
accorded a specially protected status (Hawksworth & Greuter,
1989; Hawksworth, 1991).
Resources. Ainsworth (1963) foresaw that computers would
play a major role in handling the data on fungi. The vast
numbers of species will necessitate the computerization of
comparative descriptive data (including illustrations), and not
merely names and specimen details. Some of the largest
mycological herbaria, notably that of IMI and the u.s.
National Fungus Collection, have now installed computer
systems, but find the task of keying in all past acquisitions
daunting. Rapid progress is taking place in the handling and
exchange of biosystematic information across biology as a
whole (Allkin & Bisby, 1984). Mycologists can learn from
what has already been achieved, and also contribute to the
future development of such systems (for example through the
IUBS Taxonomic Databases Working Group) to enable them
to be as widely compatible as possible.
Presidential address
The rapidly expanding CD-ROM technology, already in
use as a medium for the CAB ABSTRACTS bibliographic
database, and CD-STRAINS (Hitachi Ltd) for culture collection
information, can be expected to emerge as an appropriate
medium for the storage and dissemination of systematic
information. In time, particularly when graphic and picture-
storage capabilities improve and costs fall, CD-ROM discs
could become a more effective outlet than hard copy for
descriptive information.
Storage space for collections will also increasingly become
an issue. Dried preserved specimens provide the reference
points for the application of names and are the scientific basis
of verifiable information on distributions, substrata and hosts
(Hawksworth & Mound, 1991). Now that DNA can be
replicated from such material (Bruns et 141., 1989), and even a
single ascospore (Lee & Taylor, 1990), by means of the
polymerase chain reaction (PCR; Innis ef 141., 1990), dried
specimens are a genetic and not only a reference resource able
to complement that in culture collections (see below). At IML
for example, about 31500 species are represented by 330000
dried reference specimens; these currently occupy 1456 m of
shelving in compactors. At an equivalent level of rep-
resentation, shelf runs of 70 km would be required for a
comparable reference collection of the world's fungi. -
Scientific. If current classification systems are based on the
characteristics of just under 5 % of the world's fungi, the level
of confidence we can have that any proposed classification will
be able to accommodate the remaining 95 %of variation must
inevitably be extremely low. Even in those fungi which are
described, it has also to be recognized that key data relevant
to the development of taxonomic hierarchies and phylogenetic
reconstruction are often lacking. For this reason, and to
increase the stability of names in higher categories, it is
prudent and scientifically more honest to endeavour to
circumscribe families and orders for clearly closely allied fungi,
but not to arrange them in additional formal groupings within
the phyla (Hawksworth, 1985).
AHempts at phylogenetic reconstruction within the fungi
are similarly inhibited by this same data lack, but further by
the inadequately known fossil record, problems of convergence
and paedomorphosis (Eriksson, 198I; Hawksworth, 1987;
Thiers, 1984), and so the difficulty of being certain which
characters are plesiomorphic. Major strides can be expected
through the application of molecular techniques, especially
RNAs (Blanz & GoHschalk, 1986) and DNAs (by PCR
see above), and these are already starting to be
Important. However, in interpreting such results the complete
lack of data on so many fungi must not be forgotten.
SIGNIFICANCE
Why is fungal biodiversity significant 7 The scant aHention
fungi have received in the biodiversity debate is due in most
cases to a lack of awareness amongst biologists of their
significance in evolution, ecosystem function, human progress,
and Gaia.
648
In evolution
The origin of land plants may not have been pOSSible without
fungi. Fungi have been found in the creeping organs and stems
of the earliest rhynioids of the late Silurian and Lower
Devonian, for example Palaeomyces asteroxyli inside Asteroxylon
rhizomes. This led to the hypothesis that such plants were the
product of an association between a green alga and a fungus
(Pirozynski & Malloch, 1975); Palaeomyces, although lacking
'arbuscules', has been considered as possibly congeneric with
the modern endomycorrhizal (VAM) genus Glomus in the
Endogonales (pirozynski & Dalpe, 1989). The possibility that
these fungi were saprobes cannot be discounted, but even if
they were they would have been crucial as biodegraders
providing material on which the earliest land plants could
grow (Taylor, 1990).
The earliest filamentous fungal hyphae occurred endo-
lithically in the much earlier Cambrian shelly faunas (Taylor,
1990), interestingly, a habitat in which some ascomycetes
lichenized with cyanobacteria occur today. As noted by von
Arx (1987) there are claims of Precambrian fungi, but the true
nature of those is less clear. There continues to be a dispute
as to whether the ascomycetes had a common ancestor with
early 'phycomycetous' fungi or arose separately from red
algae; evidence for the laHer route is substantial (Demoulin,
1985). By whatever course, asci adapted to aerial discharge
must have arisen by about 500 million years ago, before or
contemporaneous with the earliest rhinioids, as in the Silurian
the ascomycetes were evidently already diverse (Sherwood-
Pike & Gray, 1985). It has been suggested that the first may
well have been lichen-like, associated initially with cyano-
bacteria and later with green algae (Cain, 1972; Eriksson,
1981; Hawksworth, 1982b, 1988 b). Indeed Church (1921)
considered that lichens were transmigrants, developing from
the sea onto the land; this view has continued to receive some
support (Corner, 1967: p. 260). Lichens could also have had a
key role in accelerating rock breakdown as they do today
(Lawrey, 1984; Jones & Wilson, 1985; Wessels & Schoeman,
1988), and so have been instrumental in the production of
material in which the first land plants were' rooted '.
Fossil trunks of the progymnosperm Callixylon newberryi
exhibit patterns of wood-decay characteristic of hymeno-
mycetes in the Upper Devonian, and could be the first
evidence of a basidiomycete, but clamp connections were not
seen (Stubblefield ef 141., 1985). The earliest documented clamp
connections are from the Carboniferous (Dennis, 1970), but
basidiomycetes were perhaps not then common or such
extensive coal measures would never have formed. As wood-
and liHer-rotting fungi evolved, they must have made an
increasing contribution to biodegradation and the develop-
ment of soils.
If rock breakdown involving fungi and the establishment of
mycorrhizas were crucial to the evolution of the land flora
without fungi there would have been no lichens or
phytes, no vascular plants, no dinosaurs to feed on them, and
consequently no man. There might even have been no insects
as mycophagy may be primitive in the group (Crowson,
1984; Pirozynski & Hawksworth, 1988: p. 11), and the
D. L. Hawksworth
enigmatic Blochmann bodies and other organelles in insect
tissues could also be of fungal origin (Smith & Douglas, 1987).
In addition to the colonization of land, fungi have had a
major role in the subsequent radiation of other groups,
especially vascular plants, through the development of
mutualisms. VAM mycorrhizas with arbuscules were definitely
present in the Triassic (Stubblefield et aI., 1987). Ecto-
mycorrhizas developed later and enabled forests to spread
from the tropics into temperate regions with fluctuating
climates and on poor soils from the Middle Cretaceous
(Malloch et al., 1980). Mycorrhizas occur in 75-80% of our
present vascular plants (Malloch et aI., 1980; Harley & Harley,
1987). Mycophyllas (see above) also place plants with
endophytic fungi at a competitive advantage, for example
against herbivores and insect pests (Carroll, 1988), and must
similarly have influenced the course of evolution.
Lamboy (1984) and Pirozynski (1988) suggested that
horizontal gene transfer by incorporation of DNA from fungi
into vascular plant genomes may have contributed to the
development, and especially diversification of, leaves, flowers,
and fruits. Gut endosymbionts in gall-fanning (often pol-
linating) insects could have played a crucial role in this process
(Pirozynski, 1991). Atsatt (1988) even proposed that vascular
plants were a mosaic of tissues of algal and fungal origins;
, inside-out' lichens.
Intimate associations continue to be important in evolution
today, and the range of coevolutionary situations identified
which involve fungi is impressive (Pirozynski & Hawksworth,
1988). Mutualisms are especially important in this connection.
Indeed it is becoming clear that in an evolutionary context a
plant (or insect) should not be viewed as a single organism,
but rather a mutualistic association including endophytic and
mycorrhizal fungi (or gut endosymbionts). The unit subject to
natural selection, and so to the evolution of biodiversity, is
therefore the mutualism and not members of one species. As
eloquently argued by Price (1988), mutualism facilitates
adaptive radiation; fungi have had this crucial role for both
plants and animals.
In ecosystem function
Scant attention has been accorded to the role of fungi in
ecosystem function, and so in the maintenance of biodiversity
itself. Apinis (1972) drew attention to many aspects of this
topic, and Wicklow & Carroll (1981) bring together some
pertinent papers. However, broadly based integrated inter-
disciplinary studies are required to place fungi in an ecosystem
context. While the biomass involved may be considerable,
especially when below-ground and litter-inhabiting fungi are
considered, it is their functions which are crucial to ecosystem
maintenance. In addition to 75-80% of vascular plants having
mutualistic mycorrhizal fungi (see above), they have significant
roles as parasites in the natural biocontrol of other organisms,
as sources of food for a great variety of organisms (insects,
small mammals, nematodes, molluscs, etc.), mutualists in
wood-boring insects, saprobes in the breakdown and nutrient
cycling of dead plant and animal remains, carbon entrapment,
and nitrogen-fixation. The latter is often overlooked in a
649
mycological context, but is achieved through cyanobacteria
within lichen thalli; this phenomenon merits particular
attention in south-temperate and subtropical forests (Guzman
et al., 1990). Major gaps remain even in our understanding of
the interactions of phylloplane, rhizoplane and rhizosphere
fungi with crop plants, as highlighted in Grossblatt (1989).
Food webs are ultimately based on micro-organisms,
including fungi, reflecting the course of evolution (Price,
1988). Species diversity tends to be greatest amongst smaller
organisms (May, 1988), but it has been argued that much of
this variation can be functionally redundant (Norton, 1986b:
p. 122). In fungi it might be thought reasonable to assume that
all wood-decay or litter-rotting species in a site are not
necessary for that ecological function to occur effectively.
However, such 'redundancy' can also endow an ecosystem
with resilience against the loss of some species with similar
functions; such 'bootstrapping' involving fungi may be
especially important in the maintenance of soils (Perry et al.,
1989). The ability of many deciduous temperate trees to
associate with a variety of ectomycorrhizal fungi also places
them at a competitive advantage by 'bootstrapping' as
compared with unimycorrhizal species. In the British Isles
there are indications that at least 55 hymenomycetes can fonn
mycorrhizas with Quercus (Watling, 1974); 103 with Betula, 90
with Pinus sylvestris, etc. (Alexander & Watling, 1987). The
capacity to utilize a selection of species contributes to the
success of such ecosystems by providing a 'bootstrapping'
buffer to the loss of individual associates.
Conversely, in some cases fungi may be 'keystone species'
which if lost would lead to a major change in the ecosystem;
for example if the species was the only one fonning a
mycorrhiza with the dominant plant, limiting an insect that
would otherwise become a major pest, a pathogen keeping
another plant 'down', etc. While it is opportune that the
effects of ecological disturbance or change on fungi are now
being considered (Boddy et aI., 1988), in-depth ecological
studies in different ecotones remain necessary in order to
quantify the role of fungi and other microorganisms in
ecosystem function (di Castri & Younes, 1990).
Finally, fungi have a major role as indicators of ecosystem
health as monitors of disturbance of the soil, and through
lichens of above-soil ecological continuity (Harding & Rose,
1986; Thomson, 1990), and further environmental pollution
(see below). Further work can be expected to reveal new
monitoring applications.
In human progress
The just under 5 % of known fungi include some put to a great
variety and extending range of applications to further man's
progress. Their economic value world-wide has not been
quantified, but must be counted in billions of US $. Current
commercial uses include: amino acid production; antibiotics;
beers, wines, brandies and distilled alcohols; breads; biocontrol
agents; cheeses; enzymes; fennented foods; food (mushrooms
etc); flavours; food colorants; fuel (ethanol, biogas); herbi-
cides; organic acids; pesticides; preservatives; Single-celled
protein (' Quom'); soy sauce; vitamins; and waste bio-
Presidential address
degradation. Exciting new applications and products continue
to be developed from both known and new species. This is
reflected in a three-fold increase in the number of patent
strains being deposited annually at IMI since 1986.
Of particular importance at a time of public concern over
chemical pesticides is their use as biocontrol agents of fungi
causing plant diseases, of weeds, or of insect pests. They
persist in populations at low levels avoiding the need
for repeated applications. A multinational US$4'5 million
examination of their potential for locust control. launched in
1989 and co-ordinated by the International Institute of
Biological Control. is indicative of the possibilities.
Secondary metabolites have only been documented in
about 5000 of the species of known fungi (i.e. 7%). These
continue to yield new commercially important products, such
as cyclosporin used in preventing the body from rejecting
organ transplants. That many pharmaceutical companies are
actively screening tropical fungi for desired activities was to
be expected, and numerous biologically active products
clearly remain to be detected and characterized.
The range of genes in fungi far exceeds that in other
microorganisms; bacteria have much smaller genomes. The
search for useful genes has received new impetus from genetic
engineering, protoplast fusion, and now gene gun technology;
fungal genes can now be transferred into and expressed
through more rapidly growing bacterial and particularly yeast
production systems. The insertion of fungal genes for features
such as natural insecticide production directly into crop plants
can now be foreseen; most of the required technology already
exists.
The world's undescribed fungi can be viewed as a massive
potential resource which awaits realization. And a time when
the need to find new and improve sources of food in
particular is greater than at any period in its history.
In Gaia
The concept of Gaia (Lovelock, 1988; Margulis & Lovelock
1989), has focused attention on the need to consider the earth
in the total context of atmosphere, oceans, biota, and
lithosphere; it has further highlighted the major contribution
of microorganisms to the breakdown of rocks and composition
of the atmosphere. It seems impossible to quantify the
importance of fungi in major earth processes at this time, but
they must be considerable.
(i) The biomass of fungi in soils, plant parts, and rocks must
be substantial. especially in temperate forests and lichen-
dominated regions, and therefore traps significant amounts of
carbon. They may be major components of soil biomass,
substantially exceeding that of bacteria, nematodes, or
arthropods (Lynch, 1988). Lichens fix atmospheric carbon
dioxide photosynthetically (see above), while saprobic,
parasitic, mycorrhizal and other mutualistic fungi keep that
derived from decay materials in their tissues rather than in the
atmosphere.
(ii) Fungi involved in decay form parts of the nutrient
cycles of nitrogen, phosphorous, and sulphur as well as
carbon.
(iii) Gaseous methylated compounds relevant to atmos-
650
pheric composition can be produced by fungi, notably
methyl chloride (chloromethane) during wood decay (White,
1982), but also others such as trimethyl arsine (Cullen &
Reimer, 1989). High-efficiency methylation of halide ions by
fungi has been considered to probably make a substantial
contribution to the atmospheric methyl chloride burden
(Harper, 1985).
(iv) Accelerated rock weathering by microorganisms,
including fungi (especially those in lichens), removes carbon
dioxide from the atmosphere, for example through conversion
of silicates to oxalates (see above). The role of fungi in such
processes has as yet received scant attention compared with
bacteria (Krumbein, 1983).
While the role and so the effect of losses of fungi on Gaia
cannot be quantified, it is prudent to be concerned. As
eloquently portrayed by Durrell (1972: p. 210): 'The world is
as delicate and as complicated as a spider's web, if you touch
one thread, you send shudders running through all the other
threads that make up the web. But we're not just touching the
web, we're tearing great holes in it.'
This is especially true as Gaia may now be becoming
stressed to the extent that compensating mechanisms and
'bootstrapping' may not enable life as we know it to continue
indefinitely. By changing the climate through global warming'
we make every spot on earth man-made and artificial'
(McKibben, 1990: p. 54). The effects of the probable I-2C
rise in mean world temperatures by 2030 (Holdgate, 1989:
p. 29) on fungi are uncertain. However, they would ineVitably
involve: crops being challenged by fungi which they do not
now encounter (both as vegetation changes and potential
ranges of disease fungi are modified), leading to increased crop
losses; increased incidences of dermatophytic infections and
mycotoxicoses in regions where they are now rare, for
example in North America and Europe; changing seasonal
patterns of macromycete fruiting; and differential rates of
spread of vascular plants (including trees) and their mycorrhiza-
forming fungi hindering the migration of such associations
(and so their associated biotas). Such factors merit serious
consideration in endeavouring to forecast and model the
impact of climatic change on man, his crops, forests, and so
Gaia.
CONSERVAnON
The conservation of fungi has received scant attention in most
countries. This is regrettable in view of their role in ecosystem
function, and so the maintenance of biodiversity (see above),
but further because of the unexploited genetic resource they
represent. There are also ethical arguments (Norton, 1986a;
McNeely et aI., 1990). The conservation of fungi can be
effected by two complementary approaches, in situ and ex situ,
both of which deserve increased attention.
In situ
The in situ conservation of fungi is hampered by the lack of
information as to the species present in particular sites, the
length of time and labour-intensiveness of producing lists,
knowledge of the rareity of individual species, and in most
D. L. Hawksworth
651
CONCLUSIONS
Table S. Fungi in the world's culture collections
1 long (1989).
, Staines el al. (I986), allowing for synonymy and anamorphy.
Further, it is uncertain how many strains should be
preserved to adequately represent the infraspecific genetic
variability of a fungus. Indeed this can be expected to vary
markedly between groups. As poignantly emphasized by
Mason (1940: p. 116) 'it is not pOSSible to collect a species'.
Protocols for the preservation of fungi which it has hitherto
been difficult to maintain by cryopreservation are now rapidly
being improved (Smith, 1988). The potential for securing the
fungal genetic resource for future generations in the face of
habitat destruction consequently exists if isolates of the
species are obtained. In view of the scale of the problem,
however, this objective can only be realized through
international collaboration and massive additional resources.
254000
1
11500'
17
0'8
Strains maintained
Species represented
Percentage known species
Percentage total species
This consideration of the fungal dimension of biodiversity
presents numerous scientifically exciting new research oppor-
tunities, but also major organizational and structural
challenges.
(i) The fungi are the second largest group of organisms in
the world after the insects; the prediction of Fries (1825:
p. 47) is thus upheld. The task of completing a world inventory
is consequently of a greater magnitude than in any group of
microorganisms or plants, possibly apart from the viruses
(Table 6). In order to progress towards this, and to ensure that
fungi are adequately represented in herbaria and culture
collections, it will be necessary to substantially increase
international collaboration. Such collaboration of necessity
must encompass and build up national and regional centres in
the less-developed countries. It is in these that so many of the
unknown fungi live. An increased knowledge of tropical
mycology is a vital component of development (Subramanian,
1982).
(ii) The quantification of the importance of fungal bio-
diversity in both ecosystem function and to the maintenance
of Gaia poses many challenging questions. Fungi merit serious
attention in future debates on biodiversity, and global ecology;
both need to extend their horizons to encompass them.
(iii) The destruction of tropical and temperate habitats yet
not or inadequately explored for fungi, makes conservation a
key issue for mycologists. Knowledge of the functional
characteristics of the majority of fungi already described is
currently lacking, and it is not yet possible to forecast in which
habitats or ecosystems those with the greatest potential for
exploitation will be discovered. In situ and ex situ conservation
are therefore complementary priorities. Unlike many aspects
of science, the study of biodiversity, ecosystem function, and
the securing of the resource is time-limited. There can be little
doubt that 'future generations will find it blankly incom-
cases a lack of understanding of the precise ecological
requirements of species. Even in the relatively intensively
studied British Isles, only in the case of lichenized species is it
possible to have confidence that the database on which to
make judgements on rarity is reasonably adequate.
In situ conservation of fungi is therefore best effected by
ensuring the preservation of the widest range of least-
disturbed habitat types, and both lichens (see above) and
macromycetes can be of value in determining such sites
(Arnolds, 1991). The safeguarding of centres of plant diversity
throughout the world, which the International Union for the
Conservation of Nature and Natural Resources (1987) plans to
identify, would be a major step in securing the associated
fungi. However, many of the world's undescribed fungi are in
tropical forests undergoing massive reductions at this time.
Losses are estimated at over 100000 km
2
each year, more
than the area of The Netherlands and Switzerland combined
(Wilson, 1989). As a result, one quarter of the world's
biological diversity present in the mid-1980s is expected to
disappear during the next 25 years (Raven, 1988: p. 225); by
extrapolation, 376000 species of fungi will become extinct in
this period - over five times the number currently described.
In view of the uses to which many such fungi might have been
put to the benefits of man (see above) this destruction is
comparable to the throwing away of a food basket without
our even looking inside.
Conservation in situ will, however, only be effective if the
local populace reaps some short-term benefit from it (Bessinger,
1990). This necessitates the revision of world intellectual
property rights (Keystone Center, 1990; McNeely et al.. 1990)
so that, for example, they apply to fungal strains explOited
outside the country of origin.
Preserving a site may not ensure that the fungi present will
be conserved in perpetuity. For example, acid rain may have
been the major reason for the decline of many macromycetes
that has taken place in Austria, The Netherlands and Germany
during this century (Arnolds, 1991); this has major implic-
ations for the vegetation as a whole due to the potential loss
of mycorrhizal species. The effects of air pollutants on lichens
are well-documented, and the depletion this causes can be
dramatic (Nash & Wirth, 1987; Hawksworth, 1990); this is
potentially of major concern in the lichen-dominated tundra
(Hutchinson et al., 1987) with implications for soil erosion and
dependent organisms (including caribou and reindeer).
Ex situ
Culture collections are the botanical gardens and seed banks
of microbiologists. These hold about 254000 strains of fungi
Oong, 1989: p.262), but allowing for synonymy and
anamorphy, the listed names (Staines et al., 1986) only
represent about 11500 species; i.e. 17% of the known and just
under 1% of the world's estimated fungi (Table 5).
Nevertheless, such collections have a particular importance as
they provide a source of strains that it may be extremely
difficult or not cost-effective to endeavour to reisolate from
nature. A surprisingly large number of fungi are represented
by less than five strains, indicating the infrequency of their
isolation from nature.
Presidential address
652
Table 6. Comparison of the numbers of known and estimated total
species in the world of selected groups of organisms
, Hawksworth & Greuter (1989).
Wolf (1987).
3 M. R. Crosby (pers. comm.).
di Castri & Younes (1990).
Group
Vascular Plants
Bryophytes
Algae
Fungi
Bacteria
Viruses
Known
species
220000'
17000'
40000'
69000
3000'
5000'
Total
species
270000'
25000
3
60000'
1500000
30000'
130000'
Percentage
known (%)
81
68
67
5
10
4
on Antarctica; to Professor E. Muller (Zollikon, Switzerland)
for information on the numbers of vascular plants and fungi in
Switzerland; to Drs D. E. Shaw and G. Guymer Ondooroopilly,
Queensland) for Papua New Guinea data; to Professor J.
Webster (University of Exeter) for updated numbers of aquatic
hyphomycetes; and to many of my colleagues at IMI for
assistance in diverse ways, including Mrs V. Barkham, Mrs V.
]. Dring, Dr H. C. Evans, Ms G. Godwin, Dr P. M. Kirk, Dr Z.
Kozakiewicz, Dr C. Prior, Ms M. S. Rainbow, Dr B. C. Sutton,
Mrs P. A. Taylor, and Mrs C. Thatcher (who extracted the
data from the Biological Flora of the British Isles).
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plants and to the subsequent evolution of them and their
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I am grateful to Dr G. C. Ainsworth (Derby), Professor
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Dr M. B. Ellis (Southwold) for data on Wheatfen Broad; Dr
D.F. Farr (USDA, Beltsville) for information on host plants
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