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CURRENT STATE AND FUTURE PROSPECTS OF SPANISH

MYCOBIOTA KNOWLEDGE

HEYKOOP, M., LLARANDI, E. & MORENO, G.

Department of Plant Biology (Botany), University of Alcalá, 28871 Alcalá de Henares


(Madrid) Spain.
E-mail: michel.heykoop@uah.es

Running title: Spanish Mycobiota knowledge

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ABSTRACT

Taking Hawksworth’s estimate of the magnitude of fungal diversity as our


conceptual framework we will review the current state of mycobiota knowledge in
Spain. Therefore we will first go briefly through the history of Spanish mycology. We
will then analyze our current knowledge stressing that one of the problems which
difficult and limit the discovery and description of the missing fungi is the lack of
professional taxonomist-mycologists; the latter is especially worrying considering the
important decline in taxonomy teaching in Spanish universities. On the other hand we
will emphasize the need of extensive studies of many of the Mediterranean habitats
which despite being poorly-known, however, host very interesting and rare fungi.
Examples of these habitats are gypsipherous steppe areas, forests and wood of
Juniperus thurifera, sclerophyllous forests of Quercus, etc. Comments on these habitats
as well as on some striking fungi discovered in them will be given. In addition we will
analyze briefly the number of species new to science described by the mycologists of
the University of Alcalá during a period of ten years showing that very much work
remains to be done. Finally we will finish the article with some general conclusions.

INTRODUCTION

The most widely accepted hypothesis on the fungal dimension of biodiversity is


the estimate of Hawksworth [1], according to which there are 1.5 million species on
Earth. One of the key elements in arriving at the 1.5 million figure was the ratio
between fungi and vascular plants. The 6: 1 ratio which emerged from his analysis
concerned the numbers of fungi (including lichens) ocurring on all substrata in a given
area and not just the fungi present on plants. The fact that this author used the British
Isles’ data makes sense since it is, in the case of the fungi, the most intensively studied
region in the world. Several reasonable arguments show that this ratio, far from being
excessive, is a very conservative estimate. Moreover, Hawksworth [2] contributed with
new data which showed, ten years later, that his 1.5 million fungi estimate is too low.
Nevertheless, this author considered that it would be prudent to retain that number as a
working hypothesis.
All our research on fungal biodiversity fits within the conceptual framework
represented by Hawksworth’s [1,2] estimate. Therefore we have examined, in this
article, the situation of our knowledge of the Spanish mycobiota. If we accept that our
country has 5048 species of vascular plants [3] extrapolating the 6:1 ratio established by
Hawksworth would give a total of 30288 fungi. However, the number of vascular plants
in Spain has been reassessed and now stands at approximately 7000 species, which
means that the number of total fungi might be significantly higher (42000 species).
Moreover, Hawksworth [2] points out that the ratio of fungi to native vascular plants
should be revised upwards to 8.4:1. Therefore we have adopted a conservative figure of
40000 fungi (including lichens) in Spain. Currently there is no overall fungal check-list
available in Spain, but there are some partial catalogues for several taxonomic groups. If
we compare these partial Spanish catalogues with those of other better studied countries
we can draw some conclusions about the knowledge of the biota of these fungal groups
in our country. In some of them, e.g. Myxomycetes, it seems that our knowledge is
reasonable; however, in other such groups our knowledge is very poor and we are far
from knowing the 40000 species which make up the Spanish mycobiota.
To discover the unknown mycobiota Hawksworth [1] puts the accent on several
important questions: i) critical or even regional monographic studies yield remarkably

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high numbers of new species; ii) little-explored habitats are a major source of novel
fungi world-wide; and iii) one of the most important factors which limits the number of
new described fungi in the world is the number of taxonomist-mycologists available to
achieve this work; i.e., an important question is the available manpower. This question
will be discussed later. Another important question, which has to be taken into
consideration, is the need of regional monographic studies such as the “Flora
Micológica Ibérica” (FMI) project. And finally, it is absolutely essential to carry out
extensive studies of poorly-known habitats. Moreover, when adressing the question
“where are the missing fungi?” Hyde [4] answered raising a series of new suggestive
questions such as: “are the missing fungi in poorly studied countries?”, “are the missing
fungi on poorly studied hosts?”, “are the missing fungi in poorly studied habitats or
niches?” and “can host or tissue-specificity account for the missing fungi?”. Therefore,
our analysis of the current state of the Spanish mycobiota (see section “Where are we
now?”) will be divided into three subsections: i) in the first one we will concentrate on
the need for a sufficient number of taxonomist-mycologists (need for sufficient
manpower); ii) in the second one we will discuss the need of regional and national
floras; and, finally, iii) we will comment several Mediterranean plant communities
which are an example of still insufficiently known fungal habitats and a major source of
novel fungi.
But before we analyze the current state of our knowledge of the Spanish
mycobiota, in order to understand and evaluate it correctly, it is necessary to analyze
where we come from, i.e., we must go briefly through the history of the Spanish
mycology.

WHERE DO WE COME FROM? BRIEF HISTORY OF SPANISH MYCOLOGY.

Whereas other European countries (e.g. The Netherlands, United Kingdom, etc.) have a
long mycological tradition of several centuries, in Spain the interest for fungi is
relatively recent. It is not until the end of the 19th century (González Fragoso started in
1886 a series of papers concerning mycology, but without dealing with larger fungi), or
the beginning of the 20th century when the study of fungi started in our country. On the
other hand, the study of macromycetes except for some isolated records by Lázaro e
Ibiza [5-7], and the works published by Prof. Aranzadi concerning the mushrooms of
the Basque Country and Catalonia (between 1897 and 1914) did not start until nearly
1930.
As a matter of fact it is the great popular liking for mushrooms, which existed
traditionally in some Spanish regions such as Catalonia or Basque Country, which
boosted the study of fungi in Spain. So, at the end of the twenties and beginning of the
thirties some important authors emerged in Catalonia, such as Codina & Font-Quer [8]
who published one of the first important mycological work in Spain. The latter was
followed by a complete series of papers [9-13] which resulted in the current school
leaded by Prof. X. Llimona, at the University of Barcelona.
In other Spanish regions, such as for instance Central Spain, mycology had a
more irregular and recent development. After the last contributions of Prof. Lázaro e
Ibiza, it will not be until the thirties when an important mycological work arose in the
form of a PhD thesis [14] entitled "Macromycetes of the Guadarrama and North of
Spain", in which 112 species of Agaricales were reported. From then on until the end of
the sixties a huge gap arose in the study of fungi, not only in Central Spain but generally
in the whole country.

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Nevertheless, there were some exceptions, and apart from what happened in
Catalonia, some other Spanish regions yielded sporadically important contributions.
Such was the case of Prof. Losa España in Galicia, who from his Chair of Botany at the
University of Santiago layed the foundations of mycology in this region [15].
Other regions with a great traditional liking for mushrooms are the Basque
Country and Navarre. The first data on fungi in these regions are relatively old and were
reported by Prof. T. Aranzadi [e.g. 16]. Besides, these regions always have had several
mycological societies, which have done important contributions to Spanish mycology,
especially to its popularization.
In Central Spain, after a lapse of many years since the last contributions of Dr.
Guinea, a resurgence of mycology ocurred in the seventies thanks to Prof. F.D.
Calonge, whose first contribution was made in 1968 [17]. We can assert, undoubtedly,
that thanks to this author Spanish mycology has reached its present high level, and his
influence on the study of fungi in our country has been outstanding up to the present
time, being the founder of the mycological group at the Royal Botanical Garden of
Madrid. Prof. Calonge has promoted several PhD theses, among which are worth of
mentioning those of Profs. Moreno [18] and Ortega [19]. The latter boosted definitely
the study of fungi in Andalusia publishing numerous papers until present. Prof. Moreno
started his research creating a school of mycologists at the University of Alcalá, which
has studied a wide range of different fungal groups. This author has promoted numerous
Ph.D. theses, among which we must mention that of Prof. Honrubia [20] (co-promoted
with Prof. LLimona), who currently leads the mycological group of the University of
Murcia.
Finally, in the Canary Islands the start of mycology has been also very recent.
The first serious study of the mycobiota of this region was due to Prof. Beltrán [21].

WHERE ARE WE NOW? CURRENT STATE OF OUR KNOWLEDGE.

1) Need for a sufficient number of mycologists in order to complete the inventory of the
mycobiota. As pointed out by Hawksworth [1,2,22], one of the major problems, or
limiting factor, impeding the discovery and description of all the missing fungi is the
lack of professional mycologists/taxonomists. In Spain, the current situation might,
apparently, not look so bad or even look good. If we should establish a ranking of the
number of professional mycologists with tenure in academic or research institutions
then we would conclude that the Department of Plant Biology of the University of
Alcalá has the highest number of mycologists, 9 to be exact. It would be followed by
the University of Barcelona with 6 and the Royal Botanic Garden of Madrid with 5. In
Spain, as a general rule, taxonomist-mycologists have a botanical education.
Nevertheless, the Departments of Botany and Plant Biology, as well as the Botanic
Gardens, in which these mycologists work, except for rare cases, have been dominated
during many years by botanists specialized in vascular plants. It is not until the end of
the seventies when some botanists specialized in the field of criptogamic plants, and
more specifically in mycology, joined these Departments with a tenure. At present,
despite the important decline of new positions for taxonomists in general, and more
particularly for mycologists, the situation is not so worrying as, for example, in the UK
[22] where there remains now only one professional mycologist involved in
identification and taxonomic work in the entire university system. In contrast, in Spain
there are 14 universities or Public Research Organisms (PRO) localized in the
Autonomous Communities of Madrid, Catalonia, Galicia, Andalusia, Murcia, Balearics
and Canary Islands with professional mycologists working on aspects related with

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fungal taxonomy. This increase in the number of professional mycologists in
universities and PRO during the last 20-25 years correlates with a higher scientific
production in the same period. From a total of 4700 bibliographic 1 references which
contain records of Iberian fungi Spanish and Portuguese [24], and which range from
1762 until 1991, approximately 55% corresponds to the period from 1980 until 1991.
Despite this, apparently favourable, situation in which a certain number of
taxonomist-mycologists hold a tenure in Spanish universities and PRO, we will see
below that future prospects are not so promising, especially if we consider the important
decline in taxonomy teaching in our universities. Later we will come back to this
question.

2) Need for regional and national Floras. In Spain, as happens in other countries and
regions of Earth, we still lack a Mycological Flora. It is not until 1988 when the “Flora
Micológica Ibérica” (FMI) project was started. The objective of the FMI project is the
elaboration and edition of a work that enables identifying fungal species that grow and
fruit spontaneously in the Iberian Peninsula and Balearic Islands. On the Spanish side 4
universities (Autonomous Universities of Madrid and Barcelona, University of Alcalá
and University of Vigo) and 2 PRO (Royal Botanic Garden of Madrid and
Mediterranean Institute for Advanced Studies of the Balearic Islands) do work or have
worked on this project. The main results yielded so far by the FMI project are the
following monographic volumes:

• Vol 1: Aphyllophorales resupinatae non poroides, I. Acanthobasidium-Cystostereum


[25]
• Vol. 2: Myxomycetes, I. Ceratiomyxales, Echinosteliales, Liceales, Trichiales [26]
• Vol. 3: Gasteromycetes, I. Lycoperdales, Nidulariales, Phallales, Sclerodermatales,
Tulostomatales [27]
• Vol. 4: Laboulbeniales, I. Laboulbenia [28]

In addition, the project has also published a series of “Cuadernos de trabajo de FMI”
[24,29-45]. These “books” are generally compilations of the databases of the records of
several Spanish fungal groups; i.e., the check-lists upon which the monographic
volumes are based later on. So, check-lists of the following groups are already
available: Myxomycetes, Corticiaceae s.l., hidnaceous fungi, Tremellales s.l.,
Gasterales s.l., Ustilaginales, aquatic hyphomycetes, Dothideales, Helotiales p.p.,
Laboulbeniales, Agaricales (Inocybe, Psathyrella, Amanita, Paxillus, Hygrophoropsis,
Omphalotus, Chroogomphus, Gomphidius and Phylloporus). The number of records in
each of these groups is tabulated in table 1, in the column referring to Spain.
Hawksworth [2] provided a table (based on Rossman [46]) of the major groups
of fungi and estimated world species numbers, in which fungi are classified into three
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We must indicate that in these references are included not only published papers but also
communications presented at congresses and compiled from the abstract books. On the other hand this
figure is misleading and can distort the picture of the global magnitude, because of the fact that in works
like the “Bolets de Cataluyna”, each one of the illustrations which make up the work are compiled as one
bibliographic reference. Besides, we must also state that in the period ranging from the end of the 18 th
(1762) to the end of the 19th century the contributions are scarcer (14 references in the 18th and 200 in the
19th [23]). It is in 1886 when González Fragoso started with serious and important mycological studies
(144 of the 200 references of the 19th century were published from 1880-1899). Finally, we must stress
that lichenized fungi are excluded from these references. All the lichenized fungi are dealed with in
projects different from FMI. Here also a clear and detrimental “divorce” is established between
mycologists and lichenologists.

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different categories according to the degree of knowledge we have from them: i) well-
known; ii) moderately well-known; and iii) poorly known. From the groups studied, so
far, by the FMI project the following fit in the first category: Corticiaceae s.l. and the
hidnaceous fungi; the following in the second one: Gasterales, s.l., Ustilaginales,
Tremellales s.l., the aquatic hyphomycetes, Myxomycetes and Agaricales s.l.; and
finally, the following in the third category: Helotiales, other peritecioid ascomycetes
different from the Hypocreales and Xylariales (i.e. Dothideales), and insect-specific
fungi (i.e., Laboulbeniales). We must mention that the FMI project is also working on
Mucorales (also included in the third category) and that we lack data, except for some
isolated records, referring to Chytridiomycetes or Oomycetes considered as “poorly
known fungi”. On the other hand, we must also indicate that several groups of
micromycetes, such as Coelomycetes, dematiaceous and non-dematiaceous
hyphomycetes, Uredinales, etc. despite the fact they have not been compiled in check-
lists have been studied more or less profoundly by different authors. Their results are
scattered across a wide range of papers as, for example, those published by González
Fragoso between 1883 and 1927, or Urríes between 1932 and 1953, or the numerous
papers published by Dr. Guarro alone or in collaboration from the seventies till
present (for a complete list of all these works see the compilations carried out by Pando
[24,29]).

Table 1. Comparison of the results of the FMI project with the knowledge of several taxonomic groups in
other countries.

Number of Number of Number of Number of species


species in Spain species in The species in the in other country
Netherlands United Kingdom
Vascular flora 5048 [3] 1221 [3] 1623 [3]
Endemics of vascular 941 [3] 0 [3] 16 [3]
flora
Myxomycetes 276 [35] 309 [48] 356 (& 17 var.)
296 [47] (in both (species and [49]
cases species and infraspecific
infraspecific taxa)
taxa)
Corticiaceae s.l. 365 [31,32,34] 350 [50] 500 [50]
(species and (Scandinavia)
infraspecific
taxa)
Hidnaceous fungi 43 [51] 21 [52] 27 [53] 28 [52] (Norway)
26 [52]
(Switzerland)
32 [52] (Sweden)
Laboulbeniales 218 (sp. and var.)
[37]
Aquatic hyphomycetes 135 [45]
Tremelales s.l. 96 [38] 83 [54]
Helotiales p.p. 44 sp. (1 var.)
(fam. Orbiliaceae and [55]
Sclerotiniaceae)
Gasterales s.l. 185 (sp. and var.) 94 [54]
[36]
Ustilaginales 142 [44]
Agaricales: Inocybe 150 (sp. and var.) 118 [54]
[40]
Agaricales: Psathyrella 62 (3 var., 4 fo.) 96 [54]
[56]

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Agaricales: Amanita 39 sp. (7 var.) 21 [54]
[57]
Agaricales: Limacella 6 [57] 7 [54]
Agaricales: Paxillus 5 (1 var., 1 fo.) 4 [54] 5 [60]
[58]
Agaricales: 3 (1 var.) [58] 4 [54]
Hygrophoropsis
Agaricales: Omphalotus 2 [58] 1 [54]
Agaricales: 3 [58] 1 [54] 2 [60]
Chroogomphus
Agaricales: Gomphidius 4 [58] 3 [54] 4 [60]
Agaricales: Phylloporus 2 [58] 1 [54] 1 [60]
Dothideales 99 sp. (3 var.)
[59]

According to table 1, and taking into account the data globally, 1915 taxa
(species and varieties) of the groups studied so far in the “Cuadernos” of FMI have been
catalogued. If we compare this figure with our hypothesis of 40000 fungi in Spain the
conclusion is that only 4.79% of the total has been recorded. This is, of course, not
exactly true since numerous papers on other fungal groups are available, whose data
simply have not been listed and catalogued. But, even so, it seems clear that there still
remains much work to be done.
According to Tellería [61] 9867 species of fungi are known from the Iberian
Peninsula, belonging to the following groups:

• Oomycota 101
• Acrasiomycota 1
• Dictyosteliomycota 18
• Myxomycota 323
• Plasmodiophoromycota 3
• Ascomycota 3306
• Basidiomycota 4512
• Chytridiomycota 44
• Zygomycota 112
• Mitosporic fungi 1447

On the other hand, this author estimates that in the Iberian Peninsula (including
Portugal) there must be a total of 20790 fungal species, which would mean, according
to her, that we know approximately 47% of the total. Finally, she concludes that
according to her data we must be moderately optimistic in relation to the state of the
knowledge of the Iberian mycobiota. We, as we will show, are not so optimistic and
consider that we are far from knowing well the Spanish mycobiota. First of all, we have
assumed a number of species significantly higher: 40000 species (and probably there
might be more). This implies that we only know, approximately, 25% of the total.
Considering the partial data of table 1 we may conclude that the knowledge of
the diversity of some fungal groups in Spain (hidnaceous fungi, Tremellales s.l.,
Gasterales s.l., Amanita), is relatively good if compared with that of other European
countries better studied from a mycobiotic point of view. Other groups (Corticiaceae
s.l., Myxomycetes), despite the fact we have more or less extensive catalogues, are
probably not sufficiently well-known. Finally, there are several groups in which no
comparisons are possible: these are the cases of Laboulbeniales, Dothideales or

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Helotiales p.p.; curiously all of them are included in the category of poorly known
fungi.
Let us examine more in detail the case of two groups considered by Hawksworth
[2] as moderately well-known: i) a genus of Agaricales s.l., such as Psathyrella, which
has been studied to some extent by us and ii) the rust fungi or Uredinales. The genus
Psathyrella, though included in Agaricales (moderately well-known), is, nevertheless,
poorly known not only in Spain but also worldwide.

Table 2. Most frequently reported species of Psathyrella in Spain [56]

Species Number of records in Spain Percentage of the total


P. candolleana s.l. 290 20,6%
P. lacrymabunda 199 14,1%
P. piluliformis 131 9,3%
P. conopilus 79 5,6%
P. gracilis 74 5,2%
P. spadiceogrisea s.l 63 4,5%
P. melanthina 42 3%
P. ammophila 42 3%
P. hirta 32 2,3%
Total of the 9 most frequently reported species 952 67,6 %
Remainder: 52 species 457 32,4 %

Data of table 2 show what we have commented before: the genus is poorly
known, and the 9 most frequently reported species are probably the easiest to recognize,
since it is even possible, in many cases, to identify them macroscopically. Therefore
they use to be reported frequently in field work check-lists and catalogues, and often by
amateur taxonomists. In contrast, the rest of the species of this genus are hardly reported
when surveys or site inventories are carried out and often not even collected.

Table 3. Distribution of the genus Psathyrella in the Spanish Autonomous Communities [56]

Autonomous Community Number of records


Navarre 335
Catalonia 302
Galicia 169
Andalusia 141
Basque Country 105
Balearics 65
Madrid 57
Castile-La Mancha 53
Castile and León 47
Asturias 21
Aragon 15
Murcia 8
Valencia 5
Extremadura 3
Cantabria 1
La Rioja 1

Table 3 shows clearly, as mentioned before, the longer mycological tradition in


Communities such as Catalonia, Basque Country and Navarre. It must be emphasi-
zed that the 53 records in Castile-La Mancha correspond almost entirely (41 records) to
the paper of Heykoop & Esteve-Raventós [62]. In addition, not less than 10 Spanish
provinces (from a total of 50), representing a large surface of the country, lack any
record of this genus which, once more, proves clearly the need for more research. The

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same situation happens in other fungal groups in Spain. This type of data shows two
things: i) that, generally, data referring to the distribution of fungi, often, do not reflect
the real distribution of these organisms but that of the mycologists who study them; ii)
that extensive exploration of all the Spanish regions, in order to discover the unknown
mycobiota, is absolutely a priority.

Table 4. Comparison of the diversity of Psathyrella and the vascular flora in Spain, The Netherlands and
The United States (flora data after Groombridge [3])

Number of Number of Number Ratio of


Psathyrella vascular plant of ende- Psathyrella species:vascular plants
species species mics

Spain 61 [56] 5048 941 1:82


The Netherlands 96 [54] 1221 0 1:12
The United States 400 [63] 18956 4036 1:47

According to Hawksworth [2], in general, macromycetes are less host-specific


than micromycetes, but the numbers in a particular area compared to plants are pertinent
to overall extrapolations. Of course it would make no sense if we would assume that the
ratio Psathyrella:vascular plant in The Netherlands the highest is indicative of the
exact number there should be in Spain (see table 4). However, what is certainly clear is
that The Netherlands has a series of habitats, all of them present in Spain especially in
the northern Eurosiberian region, thus at least we might assume that all the species
found there should probably also occur in Spain. This would give an increase of more
than a 50% in the number of Psathyrella in Spain. In addition, though low, some host-
specificity in this genus does indeed exist. So, for example, Psathyrella ammophila only
fruits on Ammophila arenaria; Psathyrella halophila only on Cladium mariscus; P.
murcida and P. phegophila mainly in woods of Fagus; P. populina on dead wood of
Populus; P. typhae on culms of Typha latifolia; P. magnispora on graminae (Koeleria
vallesiana); etc. This implies that altogether the species of Psathyrella contribute to the
macromycete:plant ratio both in the Netherlands and in Spain. Obviously the higher
number of vascular plants and, especially, of endemics in Spain implies a higher
ecological diversity which, undoubtedly, implies a higher number of possible habitats.
Therefore it is not unreasonable to assume that the number of Psathyrella in Spain
should be significantly higher. On the other hand, the dutch figure is certainly more
accurate than the Spanish one as it is the result of 40 years of research by Dr. Kits van
Waveren, which culminated in his authoritative monography [64-66]. Moreover, after
his work very little new species or new records of Psathyrella have been published in
the Netherlands. In contrast, in Spain the tendency is the opposite [56,62,67-70]. The
genus has been insufficiently studied and the number of new species is increasing.
Furthermore, the unidentified material kept in our Herbarium (AH) points out to
numerous new species awaiting formal description. Finally, as already mentioned,
critical studies in different genera [4] also yield novelties. So, after a revision [70] we
now know that many of the records of Psathyrella fusca correspond in fact to P.
phegophila, new to Spain. The same happens with P. maculata, reported occasionaly as
P. populina. Heykoop & al. [70] in a critical study, not yet published, report 6 new
species to the Spanish mycobiota, which together with 2 species new to science
published recently by Esteve-Raventós & Villarreal [71] and Arenal & al. [72] give us,
so far, a total of 69 species of Psathyrella in our country. The overall conclusion is that
the data available on Psathyrella are highly insufficient.

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The case of rust fungi is also remarkable. In the UK approximately 300 species
are known [73], which give a rust fungi:plant ratio of 1:7. The rust fungi, in contrast to
Psathyrella, are very host-specific as they are obligate biotrophic parasites of vascular
plants. Many of them need two different hosts to be able to complete their life cycle.
Therefore in this case, in contrast to what happens with macromycetes, we might
extrapolate that ratio to our country. In that case, with a 1:7 ratio there might be as much
as 1000 rust fungi in Spain. Although we still lack a check-list for this group, numerous
records, scattered across literature, are available. However, the number of different
species reported does not seem to be close to 1000 (especially taking into account the
high number of synonyms published); so the rust fungi might account for a significant
part of the hidden Spanish mycobiota. On the other hand, if we compare the number of
smut fungi (672) with that of rust fungi (6929) worldwide [74] we can see that the ratio
between both is approximately 1:10; i.e. for each smut fungus there are ten rust fungi. If
we extrapolate this to Spain that would imply that for the 142 known species of smut
fungi [38] there “should” be 1420 rust fungi: even more than our estimate!
Although elaboration of Floras and Monographies is essential, it is absolutely
necessary to discover and describe fungal diversity before it disappears. Hawksworth
[1,2] stressed the need for extensive and long-term studies of insufficiently or
endangered habitats. Although more frequent in the tropics, the destruction of habitats
in our country is far from negligible. Current town plannings are often harmful to many
natural areas; in addition, the environmental conservation policies very rarely are
concerned about threatened fungi or their specific habitats. So it is no exaggeration if
we state that many of the fungi of our country might disappear before they are
discovered.
Therefore, the exploration of endangered habitats (such as the Mediterranean
ones in Spain) as well as extensive areas with little or no records at all must be a priority
(compilation of databases and check-lists within the FMI project has proved that there
are taxonomic groups which lack any record in several regions of Spain).
Several (Mediterranean) habitats of the Iberian Peninsula poorly known
according to the results of our research during the last twenty years, but, nevertheless,
rich in interesting and rare fungi will be commented below. Besides, some of the latter
are represented in table 5 showing some of the species new to science as well as new to
Spain found in them.

Table 5. (Mediterranean) habitats of the Iberian Peninsula, insufficiently studied, in which the
mycologists of the University of Alcalá have described rare or interesting new species during the last
years.

Habitat New species described New record to Spain


Forests and wood of Hyphoderma sabinicum Manjón & Lenzitopsis oxycedri Malenç. &
Juniperus thurifera Moreno [75] Bertault [80]
Trametes junipericola Manjón, Antrodia juniperina (Murrill)
Moreno & Ryvarden [76] Niemelä & Ryvarden [80]
Hamatocanthoscypha rotundispora Mycenella margaritifera (Maire)
Raitv. & Galán [77] Maas G. [81]
Xeromphalina junipericola G. Actidium nitidum (Ell.) Zogg [82]
Moreno & Heykoop [78]
Marasmiellus phaeomarasmioides
G. Moreno, Heykoop, Esteve-Rav. &
Horak [79]

Table 5. (cont.)

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Habitat New species described New record to Spain
Gypsipherous steppe areas Simocybe iberica G. Moreno & Galeropsis desertorum var.
Esteve-Raventós [83] bispora (Vasil’kov) Moreno,
Tulostoma pseudopulchellum G. Heykoop & Illana [86]
Moreno, Altés & Wright [84] Coprinus xerophilus Bogart [87]
Phaeomarasmius gypsophilus Hohenbuehelia chevallieri (Pat.)
Esteve-Rav., Villarreal, Heykoop & Pegler [88]
Horak [85] Dictyocephalus attenuatus (Peck)
Long & Plunkett [89]
Eucalyptus forests Harknessia spermatoidea Galán, G. Descolea reophylla (Bertault &
Moreno & Sutton [90] Malenç.) Malenç. [93]
Polydesmia turbinata Raitv. & Galán
[91]
Enteridium rubiginosum Gracia,
Illana & G. Moreno [92]
Descolea maculata var. occidentalis
Esteve-Rav., C. Sánchez, Villarreal
& Barrasa [93]
Scirpus holoschoenus Mycena scirpicola Villarreal, Mycena bulbosa (Cejp) Kühn.
Heykoop, Esteve-Rav. & Maas G. [96]
[94] Pholiota graminis (Quél.) Singer
Mycena hepatica Villarreal, [96]
Heykoop & Maas G. [95]
Mycena vicina Villarreal, Heykoop
& Maas G. [95]
Sclerophyllous forests of Lachnum tamajonicum Raitv. & Psathyrella dunensis Kits van
Quercus Galán [77] Wav. [62]
(especially Q. ilex ssp. ballota, Dasyscyphella patuloides Raitv. & Clitocybe collina (Velen.) Klan
Q. faginea and Q. suber)* Galán [77] [96]
Dasyscyphella inmutabilis Galán, Crepidotus subverrucisporus Pilát
Ortega & G. Moreno [97] [96]
Protounguicularia brevicapitata Galerina clavus Romagn. [96]
Raitv. & Galán [98] Inocybe pseudoasterospora Kühn.
Valseutipella multicollis Checa, G. & Boursier [96]
Moreno & Barr. [99] Volvariella caesiotincta Orton
Odonticium monfragüense Blanco, [96]
G. Moreno & Manjón [100] Phanerochaete jose-ferreirae
Lachnum cyanoparaphysatum Raitv. (Reid) Reid [100]
& Galán [101] Boletus queletii var. discolor
Graddonidiscus hispanicus Raitv. & (Quél.) Alessio [121]
Galán [102] Boletus roseoalbidus (Alessio &
Urceolella magnifica Raitv. & Galán Littini) G. Moreno & Heykoop
[103] [122]
Urceolella triseptata Raitv. & Galán Boletus permagnificus Pöder
[103] [123]
Comatricha meandrispora Castillo, Hemimycena mauretanica
G. Moreno & Illana [104] (Maire) Singer [124]
Graddonidicus hispanicus var. Agaricus lanipes (Möller &
fornicus Raitv. & Galán [105] Schäffer) Singer [124]
Luciotrichus lasioboloides R. Galán Omphalina vesuviana (Brig.)
& Raitv. [106] Bellú & Lazzari [124]
Mycena quercophila Esteve-Rav. & Lactarius atlanticus Bon [125]
Villarreal [107] Russula subazurea Bon [125]
Inocybe subtrivialis Esteve-Rav., Macrolepiota phaeodisca Bellú
Villarreal & Heykoop [108] [125]
Mycena gladiocystis Esteve-Rav. & Mycena atropapillata Kühn. &
A. Ortega [109] Maire [125]

Table 5. (cont.)

11
Habitat New species described New record to Spain
Mycena conicoalba Villarreal & Inocybe brevicystis Métrod ex
Esteve-Rav. [110] Kuyper [126]
Omphalina farinolens G. Moreno & Inocybe brunneotomentosa
Esteve-Rav. [111] Huijsm. [126]
Lactarius zugazae G. Moreno, Inocybe pelargonium Kühn. [126]
Montoya, Bandala & Heykoop [112] Inocybe tjallingiorum Kuyper
Mycena roseoquercina Villarreal & [126]
Esteve-Rav. [113] Entoloma griseoluridum (Kühn.)
Mycena bertaultiana G. Moreno & Moser [127]
Heykoop [114] Peniophora lilacea Bourdot &
Mycena rubroquercina G. Moreno & Galzin [128]
Heykoop [114] Phlebia bresadolae Parm. [128]
Lepista panaeolus var. paxilloides Phlebia nitidula (P. Karsten) Ryv.
Esteve-Rav. & Villarreal [115] [128]
Inocybe ortegae Esteve-Rav. [116]
Gymnopus castaneus Villarreal,
Heykoop & Esteve-Rav. [117]
Marasmiellus virgatocutis Robich,
Esteve-rav. & G. Moreno [118]
Gymnopus pubipes Antonín, A.
Ortega & Esteve-Rav. [119]
Hemimycena sp. nov. [120]
Palm trees Epithele canariensis (Manjón & G.
Moreno) Hjortstam, Manjón & G.
Moreno [129]
Didymium marineri G. Moreno,
Heykoop & Illana [130]
Cistus brushes Mollisina minutissima Raitv. & Teichospora trabicola Fuckel [82]
Galán [103] Russula cistoadelpha Moser &
Aleurodiscus dextrinodeocerussatus Trimbach [124]
Manjón, Blanco & G. Moreno [131] Entoloma cistophilum Trimbach
Elaphomyces trappei Galán & G. [125]
Moreno [132] Cortinarius belleri Moser [125]
Ciboria cistophila Galán, Raitv. & J.
T. Palmer [133]
Dermocybe cistoadelpha G. Moreno,
Pöder, Kirchmair, Esteve-Raventós
& Heykoop [134]
Mycena cistophila G. Moreno &
Heykoop [135]
Rectipilus cistophilus Esteve-Rav. &
Vila [136]
Inocybe rocabrunae Esteve-Rav. &
Vila [137]
Inocybe cistobulbipes Esteve-Rav. &
Vila [137]
Cladodes of Opuntia** Physarum spectabile Nannenga-
Bremekamp, Lado & G. Moreno
[138]
Badhamia grandispora Illana & G.
Moreno [139]
Didymium subreticulosporum Oltra,
G. Moreno & Illana [140]

Table 5. (cont.)

12
Habitat New species described New record to Spain
Dung of several animals Pyxidiophora fimbriata Barrasa & G. Psilocybe liniformans Guzmán &
Moreno [141] Bas [149]
Pyxidiophora crenata Barrasa & G. Preussia fleischhakii (Auersw.)
Moreno [142] Cain [150]
Strattonia dissimilis Barrasa, Solans
& G. Moreno [143]
Aphysiotroma stercorarium
Barrasa, Martínez & G. Moreno
[144]
Sordaria elongatispora Barrasa,
Lundqvist & G. Moreno [145]
Stropharia dorsipora Esteve-Rav. &
Barrasa [146]
Glabrocyphella stercoraria Barrasa,
Esteve-Rav. & C. Sánchez [147]
Bolbitius elegans E. Horak, G.
Moreno, A. Ortega & Esteve-Rav.
[148]
Sphagnum and other mosses Agrocybe setulosa G. Moreno & Hygrocybe coccineocrenata (P.D.
Barrasa [151] Orton) Moser [18]
Omphalina subglobispora G. Entoloma cuspidiferum Kühn. &
Moreno, Heykoop & Horak [152] Romagn. [18]
Cortinarius sarcoflammeus Esteve- Hygrocybe substrangulata (P.D.
Rav., Keller & Ortega [153] Orton) Moser [154]
Cortinarius bataillei (Favre ex
Moser) Hφ iland [154]
Chromocyphella muscicola (Fr.)
Donk [155]
Wood of Abies pinsapo Protounguicularia monoseptata
Galán & Raitv. [98]
Protounguicularia variepilosa Galán
& Raitv. [98]
Chromocyphella pinsapinea G.
Moreno, Ortega & Honrubia [156]
Laeticorticium pinsapineum G.
Moreno, Manjón & Hjortstam [157]
Hyphoderma galactinum Manjón, G.
Moreno & Hjortstam [158]
Hyphoderma malençonii (Manjón &
G. Moreno) Manjón, G. Moreno &
Hjortstam [158]
*
This type of habitat also occurs in other regions of the planet where, like in Spain, it is a major source of
novelty. So, mycologists of the University of Alcalá have described also numerous species new to science
in other regions (Quercus agrifoliae in Mexico), e.g. Torrendiella quintocentenaria Galán & Palmer
[159], Incrupila isabellina Raitv. & Galán, I. calcea Raitv. & Galán, Lachnum glutinosum Raitv. &
Galán, Lambertella palmeri Raitv. & Galán, Pithyella hispida Raitv. & Galán, Crocicreas sulphurea
Raitv. & Galán [105].
**
Although in Spain we have only found new species of Myxomycetes, cladodes and remains of Opuntia
represent a very selective substratum which hosts specialized fungi of other groups. So, Galán & Raitviir
[160] described Psilachnum opuntiae (an ascomycete) on cladodes of Opuntia in Mexico.

Comments on some of these habitats and species.


Sclerophyllous forests of Quercus: The mediterranean ecosystems are dominated by
evergreen shrubs and sclerophyllous trees represented, among other plants, by Quercus
spp. such as Q. ilex ssp. ballota, Q. ilex ssp. ilex, Q. suber, Q. pyrenaica, Q. coccifera
and Q. faginea, and their associated shrub communities composed by Cistus spp. (e.g.
C. ladanifer, C. laurifolius, C. populifolius, C. monspeliensis and C. albidus) as well as

13
Helianthemum spp. All these plant communities are mycologically very interesting as
they host a high diversity of fungi. The mycobiota of these areas, however, remains
poorly known and many new species are being discovered, especially during the last 10-
15 years. In addition to all the new species described by the mycologists of the
University of Alcalá (see table 5), other Spanish mycologists have also described very
interesting new species in these habitats such as, for example, Lactarius
mediterraneensis Llistosella & Bellù [161], collected in Catalonia and Italy and which
has been found later by us in the holm oak forests of Central Spain [162]. Recently,
Collado & Peláez [163], in a exhaustive study on endophytic fungi growing in Quercus
ilex and Q. faginea, reported a high number of unidentified species (79% of the total,
and the majority belonging to ascomycete groups considered as “poorly known” [2]),
many of them being new to science (e.g. Gnomonia sp., Discosia sp., Ciliophora sp.,
etc.); from the remaining 21% several are new records to Spain [Truncatella angustata
(Pers. ex Link) Hughes or Sporormiella pilosella (Cain) Ahmed & Cain]. This
represents additional evidence supporting the statement that sclerophyllous Quercus
forests are a major source of novelty. On the other hand, in neighbouring countries
which host the same habitats, new species are regularly being described; such is the
case, for example, of Italy in which Boletus comptus Simonini [164] has been described;
or France which yielded the new species Cortinarius rioussetorum Bidaud, Moënne-
Loccoz & Reumaux [165], taxa which have both to be looked for in Spain as members
of its mycobiota. As a matter of fact, B. comptus very much resembles a new species
collected by us [120], but which due to the lack of data could not be described formally;
moreover, after its first collection it was never found again. It is striking that we still
discover large new macromycetes in these Mediterranean Quercus forests, some of
them being very abundant. Such is the case in addition to Lactarius mediterraneensis,
mentioned before of Lactarius zugazae, describerd by Moreno & al. [112]. Apart from
hosting an important number of ectomycorrhizal fungi, numerous saprothrophic fungi
also occur in Quercus forests. The leaves of the Quercus trees are accumulated during
several years on the forest soil before being decayed by different folicolous fungi. These
foliose remains constitute a very interesting substratum in order to focus on the study of
many saprotrophic fungi such as Helotiales and Agaricales, and very especially on
several species of Mycena. During the last years numerous species of Mycena have been
described in this habitat and, generally, in all sort of habitats in Spain. Undoubtedly the
genus Mycena is another insufficiently studied taxon in our country which certainly will
yield much more new species. So, in addition to the characteristic Mycena of Quercus
forests (see table 5), we have described new species in i) Pinus: Mycena geesteranii
Heykoop, Esteve-Rav. & Moreno [166], Mycena gilvipes Villarreal, Heykoop & Maas
G. [94]; ii) Betula: Mycena rubescens Villarreal, Esteve-Rav., Heykoop & Maas G.
[94]; iii) Corylus: Mycena rubidofusca Villarreal, Heykoop & Maas G. [95]; iv) Salix:
Mycena olivaceoflava Villarreal, Heykoop & Maas G. [95]; v) Cistus: see table 5; vi)
Scirpus: see table 5.
All these data give us an insight into the species richness of this sort of habitats,
together with some indication of the extent of our ignorance. Not only numerous
micromycetes have to be discovered but fungi with a considerable size, and relatively
abundant, remain completely unknown.

Gypsiferous steppe areas: The xerothermic plant communities which thrive on


calcareous and gypsiferous soils in the Iberian Peninsula are of great biological interest
since they are composed of very selective and endemic species. These plant
communities are represented in Central Spain, by the Arrhenathero-Stipetum

14
tenacissimae S. dominated by Stipa tenacissima and include some interesting fungi
adapted to this very particular ecosystem. Apart from the well-known and largely
studied vascular flora [167], lichen communities and a few basidiomycete taxa have
been either found, described or studied in this habitat ([168-170] -for lichenized fungi-,
and [83,84,88,89,171-173] -for basidiomycetes-). Most of the fungal taxa which grow in
this peculiar habitat are probably endemic and highly adapted to extreme environmental
conditions such as drought and saline soils; in these areas, the soil often becomes a
gypsum crust due to high evaporation, and only highly adapted plants and fungi
(including mosses and lichens) are able to colonize these ecologically rather harsh
habitats.
An example of fungus adapted to this habitat is Phaeomarasmius gypsophilus
Esteve-Rav., Villarreal, Heykoop & Horak [85], characterized by its small size and fruit
bodies completely covered by a whitish mineral crust (which imitates the soil colour).
So far it has only been found growing on gypsiferous soils, within the Acarosporetum
placodiformis-reagentis lichen association. Its period of fructification is probably short,
and needs warm and rainy winters as it happened at the time of collection. During
numerous visits to these or similar areas in previous, much colder years, this taxon was
never found. It has been hypothesized upon the possibilty that the calcareous crust
which completely covers the basidiocarp might have two alternative causes: i) the result
of excreting the excess of salts present in hyphae due to saline soils; ii) a protection
against desiccation and the high level of UV radiation in these habitats.
On the other hand, a recent study carried out by Peláez & al. [174] showed that
from 2880 samples taken from 45 individual plants corresponding to 9 plant species
from gypsum soils, a total of 152 fungal species were recovered. All plants, except one,
harboured at least one endophyte. Only eight species were isolated from more than half
of the plant species.
In summary, all these examples represent strong evidence that even in the most
harsh habitats we must look for fungi which can easily be overlooked.

Forests and wood of Juniperus thurifera: The autochtonous forests of Juniperus


thurifera (Junipereto hemisphaerico-thuriferae) are of great interest from a botanical
and mycological point of view. These forests ocurr especially on basic soils, sometimes
mixed with Quercus ilex subsp. ballota (Junipero thuriferae-Querceto rotundifoliae),
though occasionally also on acid soils. The Juniperus thurifera formations are very
interesting mycologically and, although apparently no ectomycorrhizal taxa do occur in
the latter, they are highly selective for both parasitic and saprotrophic fungi. This is due
to the fact that its wood is very resistent to decay because of its resin and the sudden
changes of moisture and temperature ocurring in these ecosystems. Altogether this has
adapted these trees against drought. Therefore, fungi which grow on this tree must be
highly specialized biochemically in order to be able to decay this extremely hard and
resistent wood. So, it is possible to find very rare species in this vegetation,
characterized by an Iberian-North African distribution. In several surveys of these areas
we have collected some very interesting species (see table 5). One such species is
Xeromphalina junipericola Moreno & Heykoop [78], which was screened for novel
bioactive compounds and proved to synthesize a product with anti-carcinogenic
properties (Pélaez, pers. comm.).

Stems and other parts of Scirpus holoschoenus: A certain number of very selective fungi
can be found on this plant. In addition to several new species of Mycena described by
us (see table 5), we have collected other species such as Mycena bulbosa which,

15
according to Maas Geesteranus [175], fruits on Juncus, Scirpus and Glyceria, i.e.
graminae and Cyperaceae; or Pholiota graminis which, according to Breitenbach &
Kränzlin [176] fruits on graminae, and Carex or Juncus, i.e. Cyperaceae. To illustrate
the interest of this habitat it must be emphasized that we have described as much as
three diferent new species of Mycena on this one plant, which apparently are host-
specific. The latter supports, once more, the estimate of Hawksworth regarding the
magnitude of fungal diversity.

Apart from the habitats mentioned in table 5, there are numerous other
(micro)habitats which, according to our results, might host many additional new
species. Some examples are the following:

• Dried stems of Thapsia villosa: Coniochaeta areolatirubra Checa, Barrasa & G.


Moreno [177]
• Stems and branches of Cytisus: Bicornispora exophiala Checa, Barrasa, M.N.
Blanco & A.T. Martínez [178]; Coniochaeta alkalivirens Checa & G. Moreno
[177]; Coniochaeta sarothamni (Schröter) Arx & Müller (reported for the first time
in Spain by Checa & Moreno [179])]
• Woody remains of Artemisia: Leptosphaeria hispanica Checa & G. Moreno [180]
• Stems of Cakile maritima: Diademosa sabulosa Checa & M.E. Barr [181]
• Branches of Jasminum fruticans: Marasmius celtibericus G. Moreno & Raitviir
[182]
• Branches of Rubus ulmifolius: Tapesia zarza Galán [183]
• Branches of Erica arborea: Crassoascus fusisporus Checa, Barrasa & Martínez
[184]
• Branches of Fraxinus: Cryptosphaeria eunomia (Fr.: Fr.) Fuckel (reported for the
first time in Spain by Checa [185]); Gloniella sardoa Sacc. & Trav. (reported for the
first time in Spain by Checa & Moreno [179])
• Hojas de Acer: Mycosphaerella latebrosa (Cooke) Schröter (reported for the first
time in Spain by Checa [185])
• Branches of leguminosae: Cainia desmazieresii Moreau & Müller ex Krug
(reported for the first time in Spain by Checa [185])
• Fruit bodies of other fungi: Didymosphaeria winteri Niessl (reported for the first
time in Spain by Checa [185])
• Under Alnus: Alnicola scolecina (Fr.) Romagn., A. suavis (Bresad.) Kühn., Mycena
alnicola A.H. Smith, Lactarius obscuratus var. radiatus Romagn. (all of them
reported for the first time in Spain by Villarreal & Heykoop [96])
• Under Salix: Alnicola salicis (Orton) Bon, Hemimycena pseudocrispula (Kühn.)
Singer (both reported for the first time in Spain by Villarreal & Heykoop [96]);
Hebeloma fusisporum Gröger & Zschieschang (reported for the first time in Spain
by Esteve-Raventós & Heykoop [186])

All these fungi seem to be clearly host-specific. For some of them it has even been
necessary to describe a new genus to accomodate the new species; such is the case of
Crassoascus and Bicornispora. On the other hand, the following new genera have been
described in other habitats: Aphysiotroma Barrasa, A.T. Martínez & G. Moreno,
Protounguicularia Raitv. & Galán, Graddonidiscus Raitv. & Galán and Luciotrichus
Galán & Raitv. Further evidence that much more work remains to be done to know the
Spanish mycobiota properly!

16
Finally, even in more common and better known habitats, such as beech woods or
pine woods (the latter very abundant in Spain), the amount of species found new to
science or new to Spain is impressive.

WHERE DO WE GO TO? FUTURE PROSPECTS

The majority of the fungi described from the Iberian Peninsula comes from
temperate regions, from which the Eurosiberian is the best known [23]. This is the
reason why the Iberian Peninsula, located mainly within the Mediterranean region,
continues, and will continue in future, to yield many novel fungi. As we have seen
above, there are several poorly known habitats which need extensive studies to discover
the high number of unknown fungi which live in them.
As stated at the beginning of this article, the increase in the number of positions
for professional mycologists started at the end of the seventies, growing progressively
till the mid of the nineties. From then on there has been a stagnation and future
prospects within the next 15-20 years foresee a dramatic decline as a consequence of the
massive retirement of these mycologists. It is unlikely that these retirements will be
alleviated with new positions because of two reasons: i) the decline in the number of
university students due to demographic reasons, who are the main parameter
utilized to provide the departments with new positions; and ii) the long-term
reorganization of the biology degree which implies a serious and progressive decline in
taxonomy teaching, whose consequence might be an irrevocable damage to botany and
mycology. Besides, professional mycologists at universities are required to undertake
increasingly also duties other than taxonomic research.
This means that on the one hand we will lack in future the necessary human
power to describe the unknown mycobiota, and on the other hand the time necessary to
achieve this task is not on our side. The latter happens not only in Spain but also in
other European countries. So, for instance, British Fungus Flora with the issue of its
volume 8 in 1998 has covered, after approximately 30 years, less than one third of the
total project. On the other hand, Flora Agaricina Neerlandica, which has published five
volumes (and which covers only Agaricales s.l.) and started in 1988, still has to achieve
an important amount of work. The work has been planned in approximately 10 volumes,
each one covering 150-200 species. This means that the project is about halfway and
should need another 15 years to finish.
Therefore, the FMI project, which has started in more unfavourable conditions
and, in addition, covers much more than the Agaricales of Flora Agaricina Neerlandica
will need much more time to accomplish. So far, the FMI project has published only 4
volumes, which only cover some of the groups of Gasterales s.l., Myxomycetes,
Laboulbeniales and resupinate non poroid Aphyllophorales, and it has been working
during 15 years! Future prospects are therefore not very promising.
To support the statement that much remains to be done in the study of Spanish
mycobiota (if necessary at all!), we will provide a table in which the number of species
new to science described from Spain by the mycologists of the University of Alcalá in
different taxonomic groups during the period 1992-2002 is tabulated. The number of
new species described each year shows clearly that very much remains to be done in the
inventory of our mycobiota.

Table 6. Number of species new to science described from Spain by the mycologists of the University of
Alcalá in the period 1992-2002

17
Year Agaricales s.l. Gasterales s.l. Myxomycetes Ascomycetes Total
s.l.
1992 3 1 2 2 8
1993 - - 1 6 7
1994 1 - - 6 (+1 var.) 7 (+1 var.)
1995 1 - - 1 2
1996 1 1 2 (+1 var.) 2 6 (+1 var)
1997 5 (+ 1 var.) - - 1 6 (+1 var.)
1998 11 (+1 var.) - - - 11 (+1 var.)
1999 8 (+2 var.) - 1 2 11 (+2 var.)
2000 6 (+1 var.) - - 1 7 (+1 var.)
2001 7 - - - 7
2002 9 1 1 - 11

During the period 1992-2002 a total of 83 species new to science (synonimyzed


taxa have been eliminated) and 7 varieties have been described. This gives us an
average rate of 7.5 new species per year (nearly one new species per mycologist per
year), with a more or less stable or slightly increasing trend during the last years. The
two taxonomic groups in which more new species have been described are Agaricales
s.l. (52) and Ascomycetes (21). These data clearly agree with the skills of the members
of the mycology group of the University of Alcalá. It is striking that even in a group
which is supposed to be well-known worldwide (Myxomycetes) 7 new species were
described during this period. Notice that we are talking about the average new species
described per year by one research group from one university. Considering that there
are 13 more research groups in Spain (though some of them with only one mycologist)
we can get an idea of how much new species (as well as new records) are being
generated each year in Spain. This brief analysis referring to this one mycology group
shows clearly that the Spanish mycobiota is far from being well-known.

CONCLUSIONS

• In Spain the magnitude of fungal diversity is estimated at 40000 species, although it


might be higher. Consequently, and according to the data available, we know about
25% of the total mycobiota.
• In Spain, during the last 20 years there has been an important increase in the
scientific production in mycology, which correlates with a significant increase in the
number of professional mycologists in universities and other institutions. However,
a period of decline in the number of positions for professional mycologists has
started. The latter together with a serious and progressive decline in taxonomy
teaching in our universities, represents a serious impediment to the inventory of the
total Spanish mycobiota.
• In Spain several habitats of the Mediterranean region (Quercus forests, Cistus
brushes, steppe areas, etc.) turned out to host many unknown and interesting fungi.
It is striking that we still discover relatively large and abundant macromycetes new
to science.
• The mycology group of the University of Alcalá yielded an average rate of 7.5 new
species described per year (nearly one new species per mycologist per year) during
the period 1992-2002. These data, referring to this one mycology group, together
with the species richness of numerous poorly known Spanish habitats, clearly
supports the statement that the Spanish mycobiota is insufficiently known.
• Destruction of habitats in the Spanish Mediterranean region is worrying. In addition,
very little atention has been paid to fungi by nature conservationists. Despite this

18
situation some advances have been achieved in the last decade [187]. Therefore, we
need environmental policies much more concerned about threatened fungi and their
specific habitats in order to guarantee the conservation of our mycobiota.
• In summary, we may conclude that we need scientific policies more sensitive to
systematic and taxonomic research, which is absolutely essential to gain accurate
knowledge of Spanish mycobiota.

REFERENCES

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conservation. Mycol. Res. 95: 641-655.
2. Hawksworth, D.L. 2001, The magnitude of fungal diversity: the 1.5 million species estimate revisited.
Mycol. Res. 105(12): 1422-1432.
3. Groombridge, B. 1992, Global Biodiversity. Status of the earth’s living resources. A report compiled
by the World Conservation Monitoring Centre. Chapman & Hall. London.
4. Hyde, K. 2001, Where are the missing fungi? Does Hong Kong have any answers? Mycol. Res.
105(12): 1514-1518.
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130. Moreno, G., Illana C. and Heykoop, M. 1989, Contributions to the study of the Myxomycetes in
Spain. I. Mycotaxon 34(2): 623-635.
131. Moreno, G., Blanco, M.N. and Manjón J.L. 1990, Mycological studies from the National Park of
Monfragüe (Extremadura, Spain). VI. Aphyllophorales. Mycotaxon 39: 351-360.
132. Moreno, G., Galán, R. and Montecchi, A. 1991, Hypogeous fungi from peninsular Spain. II
Mycotaxon 42: 201-238.
133. Galán, R., Raitviir, A. and Palmer, J. T. 1996, Ciboria cistophila sp. nov. a leaf-inhabiting
cistophilous member of the Sclerotiniaceae. Mycotaxon 59: 227-236
134. Moreno, G., Pöder, R., Kirchmair, M., Esteve-Raventós, F. & Heykoop, M. 1997, Dermocybe
cistoadelpha, a new species in the section Sanguineae from Spain. Mycotaxon 62: 239-246.

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135. Moreno, G. & Heykoop, M. 1998, Mycena cistophila, a new mediterranean species from Spain.
Österr. Z. Pilzk. 7: 29-32.
136. Esteve-Raventós, F. and Vila, J. 1999, Rectipilus cistophilus Esteve-Rav. et Vila sp. nov., un nuevo
hongo mediterráneo. Revista Catalana Micol. 22: 1-4.
137. Esteve-Raventós, F., Vila, J. & Llimona, X. 2002, Estudios sobre el género Inocybe (Cortinariales)
en los jarales de Cataluña. I. Revista Catalana de Micol. 24: 135-145.
138. Nannenga-Bremekamp, N.E., Lado, C. and Moreno, G. 1984, A new species of Physarum
(Myxomycetes) from the Canary Isles. Mycology Proceedings C 87: 91-94.
139. Illana, C., Moreno, G. and Heykoop, M. 1992, Spanish Myxomycetes. V. A new species of
Badhamia and a new variety of Physarum. Mycotaxon 45: 241-247.
140. Oltra, M., Moreno, G. and Illana, C. 1996, A rare Didymium from Spain. Mycol. Res. 101(12):
1508-1510.
141. Barrasa, J.M. and Moreno, G. 1982, Pyxidiophora badiorostris Lundq. y Pyxidiophora fimbriata sp.
nov. en España (Pyrenomycetes) Cryptogamie Mycol. 3(1): 41-49.
142. Barrasa, J.M. and Moreno, G. 1983, Adiciones al género Pyxidiophora (Pyrenomycetes).
Cryptogamie Mycol. 4(3): 251-259.
143. Barrasa, J.M., Solans, M.J. and Moreno, G. 1985, Strattonia dissimilis (Sordariales) una nueva
especies coprófila. Int. J. Mycol. Lichenol. 2(1): 75-84.
144. Barrasa, J.M., Martínez A.T. and Moreno, G. 1985, Aphysiostroma a new nonostiolate hypocrealean
fungus. Can. J. Bot. 63(12): 2439-2443.
145. Barrasa, J.M., Lundqvist, N. and Moreno, G. 1986, Notes on the genus Sordaria in Spain. Sordaria
elongatispora, a new coprophilous species. Persoonia 13: 83-88.
146. Esteve-Raventós, F. and Barrasa, J.M. 1995, Coprophilous Agaricales from Spain. II. The genus
Stropharia and Stropharia dorsipora sp. nov. Revista Iberoamericana de Micología 12: 70-72.
147. Barrasa, J.M.., Esteve-Raventós, F., Sánchez, C., Bodensteiner, P. and Agerer, R 1998,
Glabrocyphella stercoraria, a new cyphellaceous fungus from Spain. Mycol. Res. 102 (10): 1265-
1268.
148. Horak, E., Moreno, G., Ortega, A. and Esteve-Raventós, F. 2002, Bolbitius elegans, a striking new
species from Southern Spain. Persoonia 17(4): 615-623.
149. Esteve-Raventós, F.., Bandala, V., Montoya, L. and Rubio E. 2002, Psilocybe liniformans, a new
record for theIberian Mycological catalogue. Bol. Soc. Micol. Madrid 26: 177-181.
150. Barrasa, J.M. and Checa, J. 1991, Dothideales del Parque Natural de Monfragüe (Cáceres). I. Bol.
Soc. Micol. Madrid 15: 91-102.
151. Moreno, G. and Barrasa, J.M. 1984, Agrocybe setulosa sp. nov. en España (Bolbitiaceae, Agaricales)
Cryptogamie Mycol. 5: 101-107.
152. Moreno, G., Heykoop, M. and. Horak, E 2001, A new muscicolous Omphalina with globose
basidiospores from Spain. Mycotaxon 77: 365-370.
153. Esteve-Raventós, F., Keller, G. and Ortega, A. 2001, Cortinarius sarcoflammeus sp. nov., a new
species of subgenus Dermocybe (Agaricales) growing in Sphagnum bogs. Plant Syst. Evol. 228:
219-227.
154. Heykoop, M., Esteve-Raventós, F. and Moreno, G. 1992, Algunos Agaricales interesantes de la
provinica de Guadalajara (España peninsular). Cryptogamie Mycol. 13(4): 265-281.
155. Esteve-Raventós, F. and Villarreal, M. 2000, Adiciones al catálogo micológico de los Agaricales
Ibéricos (I). Especies raras o interesantes de la zona Centro Peninsular. Bol. Soc. Micol. Madrid
25: 197-214.
156. Moreno, G., Ortega A. and Honrubia, M. 1985, Chromocyphella pinsapinea, sp. nov.
(Crepidotaceae, Agaricales), in Spain. Bol. Soc. Micol. Castellana 10: 83-88.
157. Moreno, G. and Manjón, J.L. 1987, Nuevas aportaciones al estudio de los Aphyllophorales de la
vegetación relicta de Abies pinsapo. Bol. Soc. Micol. Madrid 11(2): 261-266
158. Hjortstam, K., Manjón, J.L. and Moreno, G. 1988, Notes on select corticiaceous fungi from Spain and
North Africa. Mycotaxon 33: 257-263.
159. Galán, R. and Palmer, T. 1993, Torrendiella quintocentenaria: a new quercicolous species from
Mexico. Mycotaxon 48: 229-237.
160. Galán, R. & Raitviir, A. 1999, Psilachnum opuntiae sp. nov. (Hyaloscyphaceae) growing on Opuntia
cladodes from Mexico. Mycotaxon 72: 163-169.
161. Llistosella, J. and Bellù, F. 1996, Lactarius mediterraneensis, a new especies from the Mediterranean
region. Mycotaxon 57: 175-186.
162. Moreno, G., Prieto, F. and Heykoop, M. 2002, Adiciones al catálogo micológico de la zona centro
peninsular. I. Bol.Soc.Micol.Madrid 26:131-145.

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163. Collado, J. and Peláez, F. 2002, Hongos endófitos de Quercus ilex L. y Quercus faginea L. en la
región central de la Península Ibérica. Bol. Soc. Micol. Madrid 26: 189-209.
164. Simonini, G. 1992, Boletus comptus sp. nov. Rivista di Micologia 35(3): 195-208.
165. Bidaud, A., Moënne-Loccoz, P. and Reumaux, P. 2000, Cortinarius rioussetorum sp. nov. et
Cortinarius luteopes Secr. ss. R. Maire 1912. Bull. FAMM., N.S., 18: 21-27.
166. Heykoop, M., Esteve-Raventós, F. and Moreno, G. 1992, Mycena geesterani sp. nov. in Peninsular
Spain. Mycotaxon 45: 301-305.
167. Izco, J. 1984, Madrid verde. Ministerio de Agricultura, Pesca y Alimentación. Comunidad de
Madrid.
168. Llimona, X. 1974, Las comunidades de líquenes de los yesos de España. Universidad de Barcelona.
169. Crespo, A., and Barreno, E. 1975, Ensayo florístico y ecológico de la vegetación liquénica de los
yesos del centro de España (Fulgensietalia desertori). Anal. Inst. Bot. Cavanilles 32 (2): 873-908.
170. Pardo, F., and Cerván, M. 1993, Líquenes de los cerros yesíferos de Madrid. Quercus 83: 8-11.
171. Altés, A., and Moreno, G. 1995, Tulostoma fimbriatum, the correct name for Tulostoma readerii.
Mycotaxon 56: 421-425.
172. Moreno, G., and Altés, A. 1992, Tulostoma simulans (Gasteromycetes), una especie generalmente
mal interpretada en España. Bol. Soc. Argent. Bot. 28 (1-4): 159-164.
173. Moreno, G., Kreisel, H. and Altés, A. 1996, Calvatia complutensis sp. nov. (Lycoperdaceae,
Gasteromycetes) from Spain. Mycotaxon 57: 155-162.
174. Peláez, F., Collado, J., Arenal, F., Basilio, A., Cabello, A., Díez Matas, M.T., García, J.B., González
Del Val, A., González, V., Gorrochategui, J., Hernández, P., Martín, I., Platas, G. & Vicente, F.
1998, Endophytic fungi from plats living on gypsum soils as a source of secondary metabolites
with antimicrobial activity. Mycol. Res. 102(6): 755-761.
175. Maas Geesteranus, R.A. 1992, Mycenas of the Northern Hemisphere. 2 Vols. Kon. Ned. Akad. Wet.
Verh. Afd. Nat. II 90.
176. Breitenbach, J. and Kränzlin, F. 1995, Pilze der Schweiz. Band 4. Blätterpilze 2. Teil.
Entolomataceae. Pluteaceae. Amanitaceae. Agaricaceae. Coprinaceae. Bolbitiaceae.
Strophariaceae. Verlag Mykologia Luzern. Luzern.
177. Checa, J., Barrasa, J.M., Moreno, G., Fort, F. and Guarro, J. 1988, The genus Coniochaeta (Sacc.)
Cooke (Coniochaetaceae, Ascomycotina) in Spain. Cryptogamie Mycol. 9 (1): 1- 34.
178. Checa, J., Barrasa, J.M., Blanco, M.N. and Martínez, A.T. 1996, Bicornispora exophiala, a new
genus and species of the Coryneliales and its black yeast anamorph. Mycol. Res. 100(4): 500-504.
179. Checa, J. and Moreno, G. 1985, Estudios sobre Pyrenomycetes y Loculoascomycetes
(Ascomycotina) II. Bol. Soc. Micol. Castellana 9: 5-14.
180. Checa, J. and Moreno, G. 1987, Leptosphaeria hispanica, a new species of the Dothideales,
Ascomycotina. Can. J. Bot. 65(10): 2096-2097.
181. Checa, J. and Barr, M.E. 1999, Pyrenomycetes sensu lato from Almería (Spain) Cryptogamie
Mycologie 20(2):79-90.
182. Moreno, G. and Raitviir A. 1999, Marasmius celtibericus (Tricholomataceae, Agaricales). A new
species from Spain. Persoonia 16(4): 541-544.
183. Galán, R. 1991, Estudios micológicos en el parque natural de Monfragüe (Extremadura, España). V.
Leotiales (=Helotiales auct.) Ascomycotina. Cryptogamie Mycol. 12 (4): 257-291.
184. Checa, J., Barrasa, J.M. and Martínez, A.T. 1993, Crassoascus, a new nonstromatic genus in the
Clypeosphaeriaceae. Mycotaxon 46: 299-305.
185. Checa, J. 1994, Pyrenomycetes s. lato de reservas naturales ibéricas. I. Bol. Soc. Micol. Madrid 19:
3-14.
186. Esteve-Raventós, F. and Heykoop, M. 1998, Studies on the genus Hebeloma (Cortinariaceae) in the
Iberian Peninsula, II. New and final observations on H. cremeopallidum, a synonym of H.
fusisporum. Doc. Mycol. 28(109-110): 17-19.
187. Llarandi, E., Moreno, G. and Heykoop, M. 2003, Hongos y Conservación. Conservación Vegetal 8:
3-6.

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