Spanish My Cob Iota

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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: [email protected] Running title: Spanish Mycobiota knowledge 1

Transcript of Spanish My Cob Iota

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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: [email protected]

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 bibliographic1 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 different categories according to the degree of knowledge we have from them: i) well-1We 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 18 th 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 19 th 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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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 species in Spain

Number of species in The Netherlands

Number of species in the

United Kingdom

Number of species in other country

Vascular flora 5048 [3] 1221 [3] 1623 [3]Endemics of vascular

flora941 [3] 0 [3] 16 [3]

Myxomycetes 276 [35]296 [47] (in both cases species and

infraspecific taxa)

309 [48](species and infraspecific

taxa)

356 (& 17 var.) [49]

Corticiaceae s.l. 365 [31,32,34] (species and infraspecific

taxa)

350 [50] 500 [50](Scandinavia)

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.

(fam. Orbiliaceae and Sclerotiniaceae)

44 sp. (1 var.) [55]

Gasterales s.l. 185 (sp. and var.) [36]

94 [54]

Ustilaginales 142 [44]Agaricales: Inocybe 150 (sp. and var.)

[40]118 [54]

Agaricales: Psathyrella 62 (3 var., 4 fo.) [56]

96 [54]

Agaricales: Amanita 39 sp. (7 var.) 21 [54]

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[57]Agaricales: Limacella 6 [57] 7 [54]Agaricales: Paxillus 5 (1 var., 1 fo.)

[58]4 [54] 5 [60]

Agaricales: Hygrophoropsis

3 (1 var.) [58] 4 [54]

Agaricales: Omphalotus 2 [58] 1 [54]Agaricales:

Chroogomphus3 [58] 1 [54] 2 [60]

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 Helotiales p.p.; curiously all of them are included in the category of poorly known fungi.

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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 totalP. 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 recordsNavarre 335

Catalonia 302Galicia 169

Andalusia 141Basque Country 105

Balearics 65Madrid 57

Castile-La Mancha 53Castile and León 47

Asturias 21Aragon 15Murcia 8

Valencia 5Extremadura 3

Cantabria 1La 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 same situation happens in other fungal groups in Spain. This type of data shows two

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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 ofPsathyrella species

Number ofvascular plantspecies

Numberof ende-mics

Ratio ofPsathyrella species:vascular plants

Spain 61 [56] 5048 941 1:82The Netherlands 96 [54] 1221 0 1:12The 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.

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

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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 SpainForests and wood of Juniperus

thuriferaHyphoderma sabinicum Manjón &

Moreno [75]Trametes junipericola Manjón,

Moreno & Ryvarden [76]Hamatocanthoscypha rotundispora

Raitv. & Galán [77]Xeromphalina junipericola G.

Moreno & Heykoop [78]Marasmiellus phaeomarasmioides

G. Moreno, Heykoop, Esteve-Rav. & Horak [79]

Lenzitopsis oxycedri Malenç. & Bertault [80]

Antrodia juniperina (Murrill) Niemelä & Ryvarden [80]

Mycenella margaritifera (Maire) Maas G. [81]

Actidium nitidum (Ell.) Zogg [82]

Table 5. (cont.)

Habitat New species described New record to SpainGypsipherous steppe areas Simocybe iberica G. Moreno &

Esteve-Raventós [83]Galeropsis desertorum var.

bispora (Vasil’kov) Moreno,

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Tulostoma pseudopulchellum G. Moreno, Altés & Wright [84]Phaeomarasmius gypsophilus

Esteve-Rav., Villarreal, Heykoop & Horak [85]

Heykoop & Illana [86]Coprinus xerophilus Bogart [87]Hohenbuehelia chevallieri (Pat.)

Pegler [88]Dictyocephalus attenuatus (Peck)

Long & Plunkett [89]Eucalyptus forests Harknessia spermatoidea Galán, G.

Moreno & Sutton [90]Polydesmia turbinata Raitv. & Galán

[91]Enteridium rubiginosum Gracia,

Illana & G. Moreno [92]Descolea maculata var. occidentalis Esteve-Rav., C. Sánchez, Villarreal

& Barrasa [93]

Descolea reophylla (Bertault & Malenç.) Malenç. [93]

Scirpus holoschoenus Mycena scirpicola Villarreal, Heykoop, Esteve-Rav. & Maas G.

[94]Mycena hepatica Villarreal, Heykoop & Maas G. [95]

Mycena vicina Villarreal, Heykoop & Maas G. [95]

Mycena bulbosa (Cejp) Kühn. [96]

Pholiota graminis (Quél.) Singer [96]

Sclerophyllous forests of Quercus

(especially Q. ilex ssp. ballota, Q. faginea and Q. suber)*

Lachnum tamajonicum Raitv. & Galán [77]

Dasyscyphella patuloides Raitv. & Galán [77]

Dasyscyphella inmutabilis Galán, Ortega & G. Moreno [97]

Protounguicularia brevicapitata Raitv. & Galán [98]

Valseutipella multicollis Checa, G. Moreno & Barr. [99]

Odonticium monfragüense Blanco, G. Moreno & Manjón [100]

Lachnum cyanoparaphysatum Raitv. & Galán [101]

Graddonidiscus hispanicus Raitv. & Galán [102]

Urceolella magnifica Raitv. & Galán [103]

Urceolella triseptata Raitv. & Galán [103]

Comatricha meandrispora Castillo, G. Moreno & Illana [104]

Graddonidicus hispanicus var. fornicus Raitv. & Galán [105]

Luciotrichus lasioboloides R. Galán& Raitv. [106]

Mycena quercophila Esteve-Rav. & Villarreal [107]

Inocybe subtrivialis Esteve-Rav., Villarreal & Heykoop [108]

Mycena gladiocystis Esteve-Rav. & A. Ortega [109]

Psathyrella dunensis Kits van Wav. [62]

Clitocybe collina (Velen.) Klan [96]

Crepidotus subverrucisporus Pilát [96]

Galerina clavus Romagn. [96]Inocybe pseudoasterospora

Kühn. & Boursier [96]Volvariella caesiotincta Orton

[96]Phanerochaete jose-ferreirae

(Reid) Reid [100]Boletus queletii var. discolor

(Quél.) Alessio [121]Boletus roseoalbidus (Alessio & Littini) G. Moreno & Heykoop

[122]Boletus permagnificus Pöder

[123]Hemimycena mauretanica

(Maire) Singer [124]Agaricus lanipes (Möller &

Schäffer) Singer [124]Omphalina vesuviana (Brig.)

Bellú & Lazzari [124]Lactarius atlanticus Bon [125]Russula subazurea Bon [125]

Macrolepiota phaeodisca Bellú [125]

Mycena atropapillata Kühn. & Maire [125]

Table 5. (cont.)

Habitat New species described New record to Spain

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Mycena conicoalba Villarreal & Esteve-Rav. [110]

Omphalina farinolens G. Moreno & Esteve-Rav. [111]

Lactarius zugazae G. Moreno, Montoya, Bandala & Heykoop [112]Mycena roseoquercina Villarreal &

Esteve-Rav. [113]Mycena bertaultiana G. Moreno &

Heykoop [114]Mycena rubroquercina G. Moreno &

Heykoop [114]Lepista panaeolus var. paxilloides

Esteve-Rav. & Villarreal [115]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]

Inocybe brevicystis Métrod ex Kuyper [126]

Inocybe brunneotomentosa Huijsm. [126]

Inocybe pelargonium Kühn. [126]Inocybe tjallingiorum Kuyper

[126]Entoloma griseoluridum (Kühn.)

Moser [127]Peniophora lilacea Bourdot &

Galzin [128]Phlebia bresadolae Parm. [128]

Phlebia nitidula (P. Karsten) Ryv. [128]

Palm trees Epithele canariensis (Manjón & G. Moreno) Hjortstam, Manjón & G.

Moreno [129]Didymium marineri G. Moreno,

Heykoop & Illana [130]Cistus brushes Mollisina minutissima Raitv. &

Galán [103]Aleurodiscus dextrinodeocerussatus Manjón, Blanco & G. Moreno [131]

Elaphomyces trappei Galán & G. Moreno [132]

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]

Teichospora trabicola Fuckel [82]Russula cistoadelpha Moser &

Trimbach [124]Entoloma cistophilum Trimbach

[125]Cortinarius belleri Moser [125]

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.)

Habitat New species described New record to SpainDung of several animals Pyxidiophora fimbriata Barrasa & G. Psilocybe liniformans Guzmán &

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Moreno [141]Pyxidiophora crenata Barrasa & G.

Moreno [142]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]

Bas [149]Preussia fleischhakii (Auersw.)

Cain [150]

Sphagnum and other mosses Agrocybe setulosa G. Moreno & Barrasa [151]

Omphalina subglobispora G. Moreno, Heykoop & Horak [152]

Cortinarius sarcoflammeus Esteve-Rav., Keller & Ortega [153]

Hygrocybe coccineocrenata (P.D. Orton) Moser [18]

Entoloma cuspidiferum Kühn. & Romagn. [18]

Hygrocybe substrangulata (P.D. Orton) Moser [154]

Cortinarius bataillei (Favre ex Moser) Hiland [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 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

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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 tenacissimae S. dominated by Stipa tenacissima and include some interesting fungi adapted to this very particular ecosystem. Apart from the well-known and largely

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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, according to Maas Geesteranus [175], fruits on Juncus, Scirpus and Glyceria, i.e. graminae and Cyperaceae; or Pholiota graminis which, according to Breitenbach &

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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!

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.

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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

Year Agaricales s.l. Gasterales s.l. Myxomycetes Ascomycetess.l.

Total

1992 3 1 2 2 8

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1993 - - 1 6 71994 1 - - 6 (+1 var.) 7 (+1 var.)1995 1 - - 1 21996 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 - - - 72002 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 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.

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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.

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32. Tellería, M.T. 1992, Bases corológicas de Flora Micológica Ibérica. Números 133-249. Cuadernos de Trabajo de Flora Micológica Ibérica, 4. Servicio de Publicaciones del CSIC.

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38. Dueñas, M. 1997, Bases corológicas de Flora Micológica Ibérica. Números 1114-1223. Cuadernos de Trabajo de Flora Micológica Ibérica, 11. Servicio de Publicaciones del CSIC. Madrid.

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48. Arnolds, E., de Vries, B. and Nannenga-Bremekamp, N.E. 1995, Overzicht van niet in de standaardlijst opgenomen Myxomycotina. Pp. 832-835. In: Arnolds, E., Th. Kuyper & M.E. Noordeloos (red.). Overzicht van de paddestoelen in Nederland. Nederlandse Mycologische Vereniging.

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50. Henrici, A. 2000, An introduction to corticioid fungi. Field Mycology 1(1): 12-19.51. Illana, C. and Blanco, M.N. 2001, Bases corológicas de Flora Micológica Ibérica. Números 1572-

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52. Marren, P. 2002, Stipitate hydnoid fungi in England. Field Mycology 3(4): 120-123.53. Dickson, G. 2000, A field key to British non-resupinate hydnoid fungi. Field Mycology 1(3): 99-104.54. Arnolds, E., Kuyper, T.W. and Noordeloos M.E. 1995 (Eds.), Overzicht van de Paddestoelen in

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55. Galán, R. 2001, Bases corológicas de Flora Micológica Ibérica. Números 1624-1676. Pp. 13-14, 101-139. In Dueñas, M. & al. Cuadernos de Trabajo de Flora Micológica Ibérica, 15. Servicio de Publicaciones del CSIC.

56. Heykoop, M. 2001, Bases corológicas de Flora Micológica Ibérica. Números 1677-1744 (Género Psathyrella). Pp. 15, 139-187. In Dueñas, M. & al. Cuadernos de Trabajo de Flora Micológica Ibérica, 15. Servicio de Publicaciones del CSIC.

57. Castro Cerceda, M.L. 1997, Bases corológicas de Flora Micológica Ibérica. Números 1224-1275. Pp. 11, 36-111. In Castro Cerceda, M.L., J. Checa & F. Pando. Cuadernos de Trabajo de Flora Micológica Ibérica, 12. Servicio de Publicaciones del CSIC.

58. Heykoop, M. & Moreno, G. 2001, Bases corológicas de Flora Micológica Ibérica. Números 1745- 1765 (Boletales lamelados). Pp. 15, 187-235 In Dueñas, M. & al. Cuadernos de Trabajo de Flora Micológica Ibérica, 15. Servicio de Publicaciones del CSIC.

59. Checa, J. 1997, Bases corológicas de Flora Micológica Ibérica. Números 1285-1411. Pp. 12-14, 112-180. In Castro Cerceda, M.L., J. Checa & F. Pando. Cuadernos de Trabajo de Flora Micológica Ibérica, 12. Servicio de Publicaciones del CSIC. Madrid.

60. Watling, R. 1970, British Fungus Flora. Agarics and Boleti. Vol. 1. Boletaceae: Gomphidiaceae: Paxillaceae. Royal Botanical Garden, Edinburgh.

61. Tellería, M.T. 2002, Riqueza fúngica de la Península Ibérica e Islas Baleares. El proyecto “Flora micológica ibérica”. Pp: 153-160. In: Pineda, F.D., J.M. de Miguel, M.A. Casado & J. Montalvo (eds.). La diversidad biológica en España. Ed. Prentice Hall.

62. Heykoop, M. and Esteve-Raventós, F. 1994, El género Psathyrella (Fr.) Quél. en España. I. (especies recolectadas en Guadalajara). Bol. Soc. Micol. Madrid 19: 37-57

63. Smith, A.H. 1972, The North American species of Psathyrella. Memoirs of the New York Botanical Garden. Vol. 24.

64. Kits van Waveren, E. 1985, The Dutch, French and British species of Psathyrella. Persoonia Suppl. Vol. 2: 1-300.

65. Kits van Waveren, E. 1987, Additions to our monograph on Psathyrella. Thirteen new species, some revised keys, comments on other recently described species, and corrections and additions to our monograph. Persoonia 13(3): 327-368.

66. Kits van Waveren, E. 1992, On five species of Psathyrella with lageniform pleurocystidia including variants with utriform pleurocystidia. Persoonia 14(4): 663-670.

67. Heykoop, M. and Moreno, G. 1998, The genus Psathyrella in spain. II. Psathyrella longicystidiata sp. nov. Mycotaxon 69: 479-486.

68. Heykoop, M. and Moreno, G. 2001, Studies in the genus Psathyrella in Spain. III. Psathyrella magnispora a new species in subsection Lutenses. Z. Mykol. 67(1): 55-62.

69. Heykoop, M. and Moreno, G. 2002, Studies in the genus Psathyrella in Spain IV. Psathyrella submicrospora sp.nov. and P. Microsporoides nom.nov. Mycotaxon 83: 425-433.

70 Heykoop, M., Moreno, G. and Llarandi, E. 2003, El género Psathyrella en España. V. Algunas especies raras o interesantes. Bol. Soc. Micol. Madrid 27 (in press).

71. Esteve-Raventós, F. and Villarreal, M. 2002, Two new species of Psathyrella. Czech Mycol. 54(1-2): 83-91.

72. Arenal, F., Villarreal, M. and Esteve-Raventós, F. 2003, Psathyrella albofloccosa, a new species of section Cystopsathyra. Mycotaxon 87: 173-177.

73. Henderson, 2000, A checklist of the Rust Fungi of the British Isles. British Mycological Society. Kew.

74. Kirk, P.M., Cannon, P.F. David, J.C. and Stalpers, J.A. (Eds.) 2001, Ainsworth and Bisby’s Dictionary of the Fungi, 9th edition. CAB International.

75. Manjón, J.L. and Moreno, G. 1983, Hyphoderma sabinicum sp. nov. (Corticiaceae) Anales Jard. Bot. Madrid 40(1): 11-14.

76. Manjón, J.L., Moreno, G. and Ryvarden, L. 1984, Trametes junipericola Manjón, Moreno & Ryvarden, sp. nov. Bol. Soc. Micol. Castellana 8: 47-50.

77. Raitviir, A. and Galán, R. 1994, Some new or interesting species of the Hyaloscyphaceae from Spain. Nova Hedwigia 58: 453-473.

78. Moreno, G. and Heykoop, M. 1996, Xeromphalina junipericola sp. nov. (Tricholomataceae, Agaricales) from Spain. Z. Mykol. 62(1): 37-41.

79. Moreno, G., Heykoop, M., Esteve-Raventós, F. and Horak E. 1997, Marasmiellus phaeomarasmioides spec. Nov. (Tricholomataceae, Agaricales) from Spain. Persoonia 16(3): 405-411.

80. Manjón, J.L. and Moreno, G. 1981, Estudios sobre Aphyllophorales 1. Fructificaciones sobre Juniperus. Anales Jard. Bot. Madrid 37(2): 407-416.

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81. Moreno, G. and Manjón, J.L. 1979, Mycena margaritifera Maire dans le centre de l’Espagne. Nouvelle espèce pour l’Europe. Doc. Mycol. 10(37-38): 85-87.

82. Checa, J. 1995, Pyrenomycetes s. lato de reservas naturales ibéricas. II. Bol. Soc. Micol. Madrid 20: 91-97.

83. Moreno, G. and Esteve-Raventós, F. 1990, Gymnopilus microsporus (Sing.) Sing. y Simocybe iberica sp. nov., en España Peninsular. Rivista di Micologia 33(3): 287-292

84. Moreno, G., Altés, A. and Wright, J. E. 1992, Tulostoma pseudopulchellum sp. nov. (Tulostomatales, Gasteromycetes) and allied species. Mycotaxon 43: 479-486.

85. Esteve-Raventós, F., Villarreal, M., Heykoop, M. and Horak E. 1998, Phaeomarasmius gypsohilus, a new species from gypsiferous plant communities in Central Spain. Mycologia 90: 151-154.

86. Moreno, G., Heykoop, M. and Illana C. 1989, Studies on Galeropsis and Gastrocybe (Bolbitiaceae, Agaricales). Mycotaxon 36 (1): 63-72.

87. Moreno, G. and Heykoop, M. 1998, Type studies in the genus Coprinus (Coprinaceae, Agaricales). Coprinus xerophilus a new record in Europe. Persoonia 17(1): 97-111.

88. Moreno, G. and Heykoop, M. 1993, Hohenbuehelia chevallieri (Pat.) Pegler (Tricholomataceae, Agaricales) en los Monegros (España). Bol. Soc. Micol. Madrid 18: 193-196.

89. Wright, J. E., Moreno, G., and Altés, A. 1993, Dictyocephalos attenuatus (Gasteromycetes, Basidiomycotina) new for Europe. Cryptogamie, Mycol. 14(2): 77-83.

90. Galán, R., Moreno, G. and Sutton B. 1986, Harknessia spermatoidea sp. nov., from Spain Trans. Br. mycol. Soc. 87: 636-640.

91. Raitviir, A. and Galán, R. 1995, The genus Polydesmia in Spain. Mycotaxon 53: 447-454.92. Gracia, E., Illana, C. and Moreno, G. 1996, Enteridium rubiginosum sp. nov., a new Myxomycetes

from Spain. Cryptogamie Mycol. 17(1): 33-38.93. Esteve-Raventós, F., Sánchez, C., Villarreal, M. and Barrasa, J.M. 1997, Il genere Descolea nella

Penisola Iberica. Rivista di Micologia 40(3): 251-260.94. Villarreal, M.., Heykoop, M., Esteve-Raventós, F. and Maas Geesteranus, R.A. 1998, Further new

species of Mycena and a new section from Spain. Persoonia 16(4): 527-535.95. Villarreal, M. Heykoop, M. and Maas Geesteranus, R.A. 1999, Further new species of Mycena from

Spain-II. Persoonia 17(2): 235-244.96. Villarreal, M. and Heykoop, M. 1997, Micobiota (Agaricales s.l.) de un transecto en el Valle del

Tiétar (Ávila y Toledo). Bol. Soc. Micol. Madrid 22: 187-217.97. Galán, R., Ortega, A. and Moreno, G. 1984, Dasycyphella inmutabilis sp. nov., en la Península Ibérica

(Helotiales, Ascomycotina) Int. J. Mycol. Lichenol. 1(3): 251-260.98. Raitviir, A. and Galán, R. 1986, A new genus of the Hyaloscyphaceae. Int. J. Mycol. Lichenol. 2 (2-

3): 221-234.99. Checa, J., Moreno, G. and Barr M. 1986, Valseutypella multicollis sp. nov. Mycotaxon 25(2): 523-

526.100. Blanco, M.N., Hjortstam, K., Manjón, J.L. and Moreno, G. 1989, Estudios micológicos en el Parque

Natural de Monfragüe (Extremadura, España). III. Aphyllophorales. Crytogamie Mycol. 10(3): 217-225.

101. Raitviir, A. and Galán, R. 1992, Lachnum cyanoparaphysatum sp. nov. A Mediterranean foliicolous species of the Hyaloscyphaceae. Rivista di Micologia 35(2): 159-164.

102. Galán, R. & Raitviir, A. (1992).-Notes on Spanish leaf-inhabiting Hyaloscyphaceae. Mycotaxon 44 (1): 31-44.

103. Raitviir, A. and Galán, R. 1993, Notes on Spanish glassy-haired Hyaloscyphaceae. Sydowia 45 (1): 34-54.

104. Castillo, A., Moreno, G. and Illana C. 1993, A new species of Comatricha with incompltely reticulated spores. Mycotaxon 46: 315-319

105. Raitviir, A. & Galán, R. 1994, First contribution to the knowledge of the Helotiales of Baja California and adjacent areas. Mycol. Res. 98 (10): 1137-1152.

106. Galán, R. and Raitviir, A. 1994, Luciotrichus lasioboloides, a new genus and a new species of the Pezizales. Czech. Mycol. 47 (4): 271-274.

107. Esteve-Raventós, F. and Villarreal, M. 1997, Mycena quercophila, a new species of Mycena section Polyadelphia growing on Quercus ilex leaves. Österr. Z. Pilzk. 6: 67-70

108. Esteve-Raventós, F., Villarreal, M. and Heykoop, M. 1997, Estudios sobre el género Inocybe (Fr.) Fr. en la Península Ibérica e Islas Baleares. III. Especies recolectadas en el Valle del Tiétar (Ávila y Toledo). Revista Catalana Micol. 20: 153-162.

109. Esteve-Raventós, F. and Ortega A. 1999, Two new agarics found in Andalucia (Peninsular Spain). Mycotaxon 71: 95-103.

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110. Villarreal, M. & Esteve-Raventós, F. 1999, Mycena conicoalba, a new corticolous species from Spain. Österr. Z. Pilzk. 8: 15-18.

111. Moreno, G. and Esteve-Raventós, F. 2000, Omphalina farinolens sp. nov., a new species from the Iberian xerophytic grasslands. Micologia 2000: 393-396.

112. Moreno, G., Heykoop, M., Montoya, L. and Bandala, V.M. 2000, Lactarius zugazae a new species from Spain. Micologia e Vegetazione Mediterranea 15(2): 91-100.

113. Villarreal, M. & Esteve-Raventós, F. 2000, Mycena roseoquercina, a new foliicolous species of Quercus ilex. Österr. Z. Pilzk 9: 23-26.

114. Moreno, G. & Heykoop, M. 2000, Due nuove specie di Mycena della vegetazione mediterranea iberica. Rivista di Micologia 43(4): 303-313.

115. Esteve-Raventós, F. and Villarreal, M. 2000, Type study of Lepista rickenii (Tricholomatales) and description of L. panaeolus var. paxilloides var. nov. Mycotaxon 76: 399-409.

116. Esteve-Raventós, F. 2001, Two new species of Inocybe (Cortinariales) from Spain, with a comparative study of some related taxa. Mycol. Res. 105(9): 1137-1143.

117. Villarreal, M., Heykoop, M. and Esteve-Raventós, F. 2002, Gymnopus castaneus, a new mediterranean species from Spain. Persoonia 17(4): 661-664.

118. Robich, G., Esteve-Raventós, F. And Moreno, G. 1994, Marasmiellus virgatocutis sp. nov. (Tricholomataceae, Agaricales). Rivista di Micologia 38(2): 141-148.

119. Ortega, A., Antonín, V. and Esteve-Raventós, F. 2003, Three interesting thermophilic taxa of Gymnopus (Basidiomycetes, Tricholomataceae): G. pubipes sp. nov., G. pubipes var. pallidopileatus var. nov and G. dryophilus var. lanipes comb. nov. Mycotaxon 85: 67-75.

120. Heykoop, M. 1993, Estudio taxonómico, corológico y ecológico de los hongos pertenecientes al orden Agaricales s.l. (Basidiomycotina) de la provincia de Guadalajara. Fac. Ciencias, Universidad de Alcalá. Tesis doctoral

121. Heykoop, M. 1995, Morphology and taxonomy of Boletus queletii var. discolor, a rare bolete resembling Boletus erythropus. Mycotaxon 56: 115-123.

122. Moreno, G., Heykoop, M., González, V. and Arenal, F. 1995, Suillus bovinoides (Blum) Bon and Boletus roseoalbidus (Alessio & Littini) comb. nov. Two interesting Mediterranean species. Doc. Mycol. 25 (98-100): 269-277.

123. Moreno, G. and Esteve-Raventós, F. 1988, Boletus aemilii Barbier, B. permagnificus Pöder and Xerocomus truncatus Singer, Snell & Dick, in Spain. Lazaroa 10: 253-258.

124. Moreno, G. and Esteve-Raventós, F. 1988, Estudios micológicos en el Parque Natural de Monfragüe (Extremadura, España). I. Agaricales. Bol. Soc. Micol. Madrid 12: 67-83

125. Moreno, G., Esteve-Raventós, F. and Illana, C. 1990, Estudios micológicos en el Parque Natural de Monfragüe y otras zonas de Extremadura (España), IV. Agaricales. Bol. Soc. Micol. Madrid 14: 115-142.

126. Heykoop, M. and Esteve-Raventós, F. 1994, El género Inocybe (Fr.) Fr., en la provincia de Guadalajara (España Peninsular). Bol. Soc. Micol. Madrid 18: 71-86.

127. Ortega, A. and Esteve-Raventós, F. 1996, Contribution to the study of the mycoflora of Andalucia (Spain) XI. Agaricales IV. Nova Hedwigia 62: 157-170.

128. Vizoso, M.T., Ortega, A. and Manjón J.L. 1991, Primera contribución al conocimiento de los Aphyllophorales s.l. de las comunidades naturales de Andalucía. Bol. Soc. Micol. Madrid 15: 153-164.

129. Manjón J.L. and Moreno, G. 1982, Cerocorticium canariensis sp. nov. (Corticiaceae) Botánica Macaronésica 10: 27-32.

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.

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.

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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.

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.

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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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