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Biodiversity and Conservation 7, 369384 (1998)

Inventorying the fungi of Mexico

Instituto de Ecologa, Apartado Postal 63, Xalapa, Veracruz 91000, Mexico

Received 18 March 1997; revised and accepted 6 June 1997 Conservation of the vegetation, analysis of the knowledge of the diversity of fungi, new biological methods and estimates of the numbers of species of fungi that grow in Mexico are discussed. The great forest destruction that is occurring and the loss of fungal diversity in the country is seriously depleting total biodiversity, mainly in the tropics. Methods to establish the number of Mexican species of fungi are proposed based on an average of: (1) the numbers of fungal obligates of vascular plants and animals and of the saprobic species; (2) extrapolating the number of the British fungi to Mexico, taking the British mycota as the best known in the world; and (3) extrapolating to the country the number of species known to occur in the State of Veracruz (Mexico), the best known in the country. The average of these three calculations yields about 200 000 species of fungi for Mexico. Of these, only 3.5% are known, despite considerable progress having been made in the inventory of the country's fungi in the last 30 years. It is concluded that it is necessary to place more emphasis on alpha taxonomy, to train more specialists and to prepare monographs of several groups of fungi. Keywords: biodiversity; Mexican fungi; estimations; species numbers.

Introduction Because of the simple method of nutrition of fungi, by absorption through their surface membranes and their powerful extracellular enzymes, they can grow in and on a wide range of substrata. Moreover, fungi have the unique ability to break down complex substrates, such as lignin, cellulose, chitin, keratin and others. These conditions place the fungi among those organisms which play leading roles in ecological processes vital for ecosystem maintenance (Subramanian, 1982; Rossman, 1994). However, fungi have received only scant attention to date, despite their potential as food sources or medical products. A large number of fungi are dependent on vascular plants and others on animals and humans; some species are even dependent on other fungi (Subramanian, 1982; May, 1991; Hawksworth, 1991, 1995a; Hawksworth and Ritchie, 1993). Because they reect perturbation, fungi can be used in ecology as indicators of vegetational or atmospherical changes (Guzman, 1994a; Hawksworth, 1995a). In Mexico there is both great diversity and long traditions relating to fungi as in other tropical countries (Subramanian, 1982). There are more than 40 species of sacred or hallucinogenic fungi in Mexico, of the 80 known in the world; many species of fungi have importance in traditional medicine, and around 300 species of edible mushrooms are consumed by the people; these last commanded considerable economic value. In 19931995, for example, Tricholoma magnivelare, Boletus edulis, Cantharellus cibarius and several species of Morchella were exported to the USA, Europe and Japan. But the fungi of many regions, especially those that grow in the tropics, are so poorly studied that there is still no clear picture of how many species live not only in Mexico, but on the planet (Hawksworth, 1993a; Harper and Hawksworth, 1994). After more than two centuries of study, most fungi in the world remain to be
0960-3115 1998 Chapman & Hall



discovered, especially in tropical inventory studies. To know the fungi is more than an exercise in curiosity; it is a necessity to understand their diversity, their role in nature, and their potential utility, before many of them disappear (Subramanian, 1982; May, 1990, 1991; Hawksworth and Colwell, 1992; Hawksworth and Ritchie, 1993; Smith et al., 1993; Harper and Hawksworth, 1994; Rossman, 1997). The main objective of this paper is to analyse our knowledge of the diversity of fungi in Mexico, the status of these studies and methods used to estimate the number that actually grow in the country. It seems that no such investigation has been attempted before for any tropical country, and no such study has been concluded in Mexico except for recent approximations (Guzman, 1995a, 1996), and some general observations on the country's total biodiversity (Williams-Linera et al., 1992; Toledo and Ordonez, 1993). Methods The present contribution is based on an exhaustive revision of all the available literature on the biodiversity of the fungi (including the Global Biodiversity Assessment, Heywood and Watson, 1995) and on the Mexican fungi reported from the last century to the present, based in part in the review of Guzman (1990), and also includes a review of all the volumes of the Bulletin of the Botanical Society of Mexico (vols 156, 19401995), Mexican Society of Mycology (vols 120, 19681985), Acta Botanica Mexicana (vols 137, 19881996), Micologa Neotropical Aplicada (vols 18, 19881995), and of the Revista Maxicana de Micologa (vols 112, 19851996), as well as other series or works (e.g. Moreno-Mart nez, 1988; God nez and Ortega, 1989; Herrera and Ulloa, 1990; Bandala et al., 1993). Most of the eld observations included are based on my experience during more than 42 years of work in Mexico and Latin America. Herbarium abbreviations are following Index Herbariorum (8th edn, 1990). The development of Mexican mycology The rst observations on Mexican fungi were made by indigenous people more than 500 years ago, as shown in their codices, stones and pottery gures. Their knowledge was so extensive and intensive that from such works it is now possible to understand the use or properties of several species of sacred, medical, poisonous or edible fungi common in Mexico (Guzman, 1992, 1997). In the second half of the last century and in the beginning of the present, European and North American travellers made personal observations on the fungi found in Mexico, or sent their collections to specialists (e.g. Kickx, 1841; Fries, 1851). But Mexican mycology only truly started between the 1940s and 1960s, with studies on relationships of yeasts or pathogenic fungi on plants and humans, and with the rediscovery of hallucinogenic mushrooms. The Mexican Society of Mycology was founded in 1965, which started a vigorous new period for Mexican mycology. Studies on diversity, however, concentrated on terrestrial species with a few exceptions of aquatic or subaquatic fungi (Cespedes and Castillo, 1982; Gonzalez and Herrera, 1993, 1995; Gonzalez, 1997). The number of species of agarics reported from Mexico provides an example of the history of the development of Mexican mycology. Between 1822 and 1971 (149 years), 424 species were recorded, but in 19721987 (15 years), 784 species were added (Bandala et al., 1988). This clearly shows the evolution of systematic studies in the country. By scrutinizing the origin of such works, we found that only 28% were contributed by Mexicans from

Inventorying the fungi of Mexico


Figure 1. Psilocybe uxpanapensis, one of the many tropical species of fungi in Mexico that are at risk of disappearance as a result of tropical forest policies. The species was the rst hallucinogenic mushroom to be found in an undisturbed tropical forest.

1822 to 1971, contrasting with 82% from 1972 to 1987. In the macromycetes, 582 species were reported by 1977 (Guzman, 1977). These gures show how variable the studies on the fungi are and how dependent they are on specialists. Scleroderma and Psilocybe are the only genera comprehensively and critically mono graphed for Mexico (Guzman, 1970, 1983a, 1995b). Monographs of Agaricus, Albatrellus, Amanita, Collybia, Crepidotus, Gymnopilus, Hypoxylon, Lactarius, Lentoramaria, Russula, Panaeolus and Phaeocollybia, and the Xylariaceae, Gomphaceae, Boletinae, telephoroid fungi, tremeloid fungi, and hydnaceous fungi, all macromycetes, and studies on the soil hyphomycete fungi and marine fungi are just starting, all of which will strengthen our knowledge of the Mexican mycobiota (Guzman and Perez-Patraca, 1972; Marmolejo et al., 1981; Medel et al., 1989; San Mart n and Rogers, 1989, 1993; Singer et al., 19901992; Perez-Silva and Herrera, 1991; San Mart n, 1992; Heredia, 1993; Bandala, 1994; Montoya, 1994; Valenzuela et al., 1994; Guzman-Davalos and Guzman, 1995; Mena-Portales et al., 1995; Cifuentes, 1996; Gonzalez, 1997). Among these studies, Gymnopilus and Phaeocollybia are good examples of increasing Mexican knowledge. Before 1986 only eight



species of Gymnopilus were known in the country, but after Guzman-Davalos and Gu zman (1995) had researched the genus, the total rose to 30, of which 15 have been described as new species. Only three species of Phaeocollybia were known in Mexico, but after the studies of Bandala (1994) 19 species are now recognized, nine of which were described as new. In some preliminary and recent studies on fungi carried out by my colleagues (Guzman et al., 1997) in the State of Veracruz, more than 900 species were reported, of which more than 40 are new to the country. In other studies carried out with my students (unpublished data), we found around 100 species of macromycetes from Veracruz, of which seven are new for Veracruz, six are new for the country and six are new species to science. These studies demonstrate that intensive investigations can increase our grasp of diversity and that much more needs to be done in Mexico. Compare for example, these gures with some from the USA: for example, Smith (1972) reported 431 species of Psathyrella of which 288 were described as new species. This contrasts with 26 species known from Mexico of which 21 are from the tropics, nine of which are only known in Mexico (Guzman et al., 1988). The major ascomycete studies in Mexico are focused in the contributions of Cifuentes, Chacon, Denison, Galan, Medel, Paden, San Mart n and Trappe, although the main collections are in ENCB, ITCV, FCMEX and XAL (Korf, 1997; Guzman, unpublished data). However, knowledge of the group is still poor, for example in the Phyllachoraceae Cannon (1997) argues that there are more than 1150 species known in the world, out of an estimated 157 800 species, but Guzman et al. (1997) only reported ve from Mexico. Whalley (1997) reported that in recent studies on Xylariaceae from Mexico, a quarter to one-third of the species were new, based on San Mart n and Rogers' studies (1989, 1993). In the case of mycorrhizal fungi, studies to date have been predominantly carried out in temperate regions, and little has been done in the tropics. Based on my approximation, about 2000 species of moulds, yeasts and parasitic fungi are reported from Mexico. These are distributed as follows: more than 1500 are obligate on plants, 140 species are saprobes in the soil and other substrata, 100 are on storage grains, 70 are in traditional beverages, 60 are parasitic in animals and humans, 30 are aquatic* and 20 occur on dung; this totals 1910 species. Certainly, the commonest species in several habitats have not yet been counted. There are about 4800 species of macromycetes, divided into 2400 ascomycetes (including 1800 lichenized species), 2200 basidiomycetes and 200 myxomycetes. This means that only around 6710 species of fungi are known in Mexico; rounded to 7000 this is too low considering the high biotic and geographical diversity of the country as discussed below. There are no more than ten institutions in Mexico concerned with systematic studies of fungi, and these have very few specialists. The reference collections of fungi in the country, critical for reliable diagnosis, comprise about 220 000 dried specimens dispersed through 20 institutions, of which ENCB and XAL have more than 60%, and FCME, IBUG, ITCV, MEXU, University of Tlaxcala and HEIMI 35% (Guzman, 1994b). Unfortunately many of these collections are not in an adequate condition for study, because they lack important information, as happens in many tropical countries, some of which have no collections at all (Harper and Hawksworth, 1994; Hawksworth et al., 1996; Hawksworth and Ritchie, 1993). Many fungi can be brought rather easily into pure culture, as stressed by Korf (1997), but there are only around 1000 strains of living cultures of fungi in no
*Gonzalez (1997) reported 22 species from marine habitats, which were not allowed for in this account.

Inventorying the fungi of Mexico


more than ten institutions in Mexico (Guzman, 1994b). Hawksworth (1995b) listed ten institutions with 279 strain names in Mexico, 31 with 6758 in the USA and 25 with 4505 in the UK. Cladistic and molecular studies on fungi are just beginning in Mexico, e.g. Villegas and Cifuentes in FCME on clavariaceous fungi, and those of Gonzalez in the Instituto de Ecolog a at Xalapa on the Rhizoctonia complex. Biochemical studies on the cell walls of fungi are being carried out by Ruiz-Herrera's group in Guanajuato (Guzman, 1990). Such mycological inventories as have been attempted in Mexico are mainly from the temperate forests, particularly the coniferous ones. However, tropical and subtropical forests would be expected to be the richest sources of mycological diversity. For example Guzman (1983a, 1995b) found 49 species of Psilocybe in Mexico, of which 39 were described from the tropics and subtropics in spite of the fact that the coniferous forests were the most explored (Fig. 1). Forest conservation and the diversity of fungi There is great need in Mexico to study the complex diversity of the indigenous fungi; it is necessary to train young students in this eld, but as this will take a considerable time, we need as a rst step a strong vegetation conservation programme. The continued denudation of forests, especially those from the tropics, for the short-term benets of wood and fuel has not only led to an erosion of soil, but also to the destruction of biodiversity (Subramanian, 1982; Kruk et al., 1988; Wilson, 1988). There is an urgent need to conserve the forests, as their continued destruction puts many species of fungi and other organisms in danger of extinction. Indeed, it is necessary to realize good management practices for the environment, because Mexico is seriously depleting its biodiversity by deforestation, especially in the tropics. More than 90% of the country's tropical forest has been destroyed during the last decades, as I have observed, and is well documented by Toledo and Ordonez (1993). But what do we need to do to conserve biological diversity in this country? We have several biological reserves or national parks (Gomez-Pompa and Dirzo, 1995), but with poor conservation programmes. Halter (1994) argued, critically, that biodiversity must be conserved not only in protected areas, but also outside of them. That is absolutely necessary, but how? The regenerative power of the forest must be protected and better understood, because when the forests are clear cut, herbaceous and shrubby growth very soon becomes dominant, and many fungi and other organisms are displaced from their natural habitats. Further, if this continues in the forest environment, the species with poor dispersal powers may never recolonize areas that become reforested (Rose and Wallace, 1974), as I have observed with Psilocybe aztecorum and other mushrooms in Mexico in some reforested Pinus forests. Also, because tropical forests usually grow on nutrient poor soils, it is very important to understand the broader implications of forest destruction. Overpopulation, habitat destruction and soil deterioration are some of the problems in the forests of Mexico; if these trends continue, it can be expected that the tropical wild forests will disappear in the future (Guzman et al., 1996, 1997). How much diversity is being lost and how fast is diversity declining? The Mexican government recently published for the rst time a list of all the species considered in danger of extinction (Diario Ocial, 1994); among these are several mushrooms I proposed for inclusion. The destruction of the original vegetation is so rapid that if one were



to nd an interesting species, it is quite possible to return to that site a few months later and nd it destroyed; this is a common occurrence in the tropical forests (Wilson, 1988, 1992). I have found several new species or new records of fungi in the Mexican tropics, such as lepiotaceous fungi, Psilocybe, Scleroderma and other fungi (Guzman, 1983a,b, 1995b; Guzman and Guzman-Davalos, 1992; Guzman et al., 1996, 1997) which are now impossible or very dicult to nd because of the almost complete destruction of some forests. We urgently need programmes to encourage the conservation of the vegetation and strengthen the protection of our nature reserves. Unfortunately, in many apparently protected areas, including some national parks and biosphere reserves, deforestation and agricultural and cattle grazing activities remain uncontrolled, contrary to the `ocial' programmes. It is denitely necessary to link economic development with conservation. The current reduction of biodiversity in Mexico seems destined to cause an economic catastrophe. The forests of the central part of the State of Veracruz (e.g. Cofre de Perote) are being destroyed so rapidly and intensively that the yield of edible mushrooms is decreasing. In 19831986 the yield of edible fungi fell from 747 to 157 kg per hectare, in 1990 it was 136 kg, in 1994 it had declined to 20 kg per hectare, and in 19951996 to only a few grams (based on unpublished calculations made with colleagues); these high rates of decline in production are alarming. In the Valley of Mexico, Mexico City, air pollution is also damaging mushroom production, most notably in the coniferous forests close to the city where, due to the direct or indirect high levels of air pollution, mushroom production is in a strong decline even where the forests still remain. I observed this years ago, and in Europe the decline of mushrooms due to acid rain has also been observed (Svrcek, 1993). Estimates of the number of fungi in Mexico It is impractical to know all the species of fungi that grow in a region, but it is possible to estimate indirectly or approximately the number of fungi in a region. We can base our consideration on available data sets and then make approximations by extrapolation. New programmes are being proposed to inventory all the organisms in the world (e.g. Systematics Agenda 2000), or at least all the species from a site (Rossman, 1994; Hawksworth et al., 1997; Hyde and Hawksworth, 1997). Substantial progress has been made recently in fungi, but a still greater emphasis is necessary in inventory programmes because of the lack of specialists. A full inventory of all the species of fungi in a site cannot be done by only a few mycologists, as already noted by Rossman (1994). Hawksworth et al. (1997) proposed 36 dierent niches and microhabitats in which to examine fungi in a tropical rain forest, and 21 dierent groups of mycological specialisms to study (although some of those are extremely complex, e.g. `mushrooms'). This will require a large team of specialists, particularly if the site is located in a tropical region. In order to inventory, in a simple way, all the fungi in a site (tropical or not), in at least 20 years, we need to consider 22 (if not more) groups of fungi: (i) (ii) (iii) (iv) (v) (vi) micromycetes micromycetes micromycetes micromycetes micromycetes micromycetes from from from from from from soil, water, dung, plant debris, cadavers, garbage,

Inventorying the fungi of Mexico (vii) (viii) (ix) (x) (xi) (xii) (xiii) (xiv) (xv) (xvi) (xvii) (xviii) (xix) (xx) (xxi) (xxii) endophytic fungi, parasitic fungi on plants, parasitic fungi on mammals, parasitic fungi on insects, parasitic fungi on other animals, endomycorrhizal fungi, tremellaceous fungi, aphyllophoraceous fungi (excluding the polyporaceous fungi), polyporaceous fungi, agarics, gasteromycetes, hypogeous fungi, cup fungi, pyrenomycetes, lichenized fungi, and myxomycetes.


We need to have at least two specialists for each group; thus 44 specialists are required, and each one will need at least one good technician. Therefore a total of 88 specialists will be necessary to study all the fungi from a particular site in Mexico. This is impossible because we do not have that many mycologists. In Mexico the number of professional systematic mycologists is no more than 20, working in a few taxonomic groups. In addition, many of those also have teaching and other duties and so cannot devote time to such tasks. Undoubtedly a complete inventory is impossible on any scale. In order to study and know all the fungi of a region, we need rst to train more systematists and also to prepare monographs of genera both clearly impracticable in the short time before much of the forests have been lost. It is necessary to separate two approaches for an inventory of fungi or other species rich groups: (1) to know all the species of fungi from a site; and (2) to estimate how many species are there. Fungi are closely associated with vascular plants because of their strict parasitism or obligate relationships with them. By estimating the number of plants, it is possible to estimate the number of obligate fungi. In Mexico there are about 30 000 species of vascular plants (Rzedowski, 1992), which implies that there are at least 30 000 species of fungi. But if we also consider how many fungi there are in Mexico based on those in or on insects, soil, lichen-formers, mosses and algae, ectomycorrhizal or non-ectomycorrhizal fungi, other saprobic fungi, and other fungi; or based on extrapolations of the known fungi from a well-studied region, we can perhaps get a better estimate of the answer to the important question, how many species are there in Mexico? A preliminary calculation of the fungi in Mexico (Guzman, 1995a, 1996) suggested 120 000140 000 or 185 000 species, respectively. I propose to follow three steps in order to estimate how many species of fungi inhabit Mexico. The rst is to calculate the approximate number of vascular plants, animals and saprobic species of fungi. As there are 30 000 vascular plants, not less than 30 000 species of fungi grow in the country. Approximately 140 000 species of animals are known in Mexico (Williams-Linera et al., 1992), of which probably 50% may be in some relationship with fungi, yielding 70 000 species of fungi associated with animals. There are probably at least another 100 000 species of fungi not in relationship with plants or



animals: those widely distributed in the soil, the water fungi, the leaf fungi, dung fungi, lichinized fungi and bark or wood fungi, among many others, considering the high diversity of the country. In this way we obtain 200 000 species of fungi as a rst calculation. A second method is to estimate the number of Mexican fungi by extrapolating from the number of fungi in the UK, because the British fungi are probably the best known in the world (Hawksworth, 1991). The UK has an area of 244 000 km2; Mexico is 8.07 times larger. If the UK has 12 000 species of fungi (or more, because more than 40 new species are described annually), there could be around 100 000 species in Mexico. It is clear that the weather in the UK is not the same as in Mexico, since tropical forests cover a third of the country, and more than 50% of the area is desert. Taking these factors into account, I think that it is reasonable to extrapolate the British data to Mexico. A third method of estimating how many species of fungi inhabit Mexico is to base the calculation on the number of species known to occur in a well-known Mexican region and to extrapolate those to the rest of the country. The state of Veracruz has a high biodiversity and is mycologically the best known in the country. For example, 186 species of polypores are known in Veracruz, compared to 85 in Morelos state, 84 in Nuevo Leon State, and 76 in Hidalgo State (Banala et al., 1993). Fungi from Veracruz have been studied since the last century, rst by European travellers who sent their specimens to speialists, and then by visits of several national and foreign mycologists (Guzman, 1990). For more than ten years INIREB and subsequently the Institute of Ecology in Veracruz has had a team of specialists collaborating in a variety of elds of fungal research, including systematics, culture and production of wild edible species (Bandala et al., 1988, 1993; Guzman, 1997; Guzman et al., 1996, 1997). In this way more than 1400 species of macromycetes have been reported from Veracruz, which probably represents 10% of the total, since between 3 and 20% of several groups of fungi are known in Mexico (e.g. 3% of thelephoraceous fungi, 3.5% of lepiotaceous fungi, and 20% of polyporaceous fungi) (estimates based on Marmolejo et al., 1981; Guzman and Guzman-Davalos, 1992; Bandala et al., 1993). In Veracruz there are consequently more than 14 000 species of macromycetes. It is more problematic to evaluate the micromycetes in Veracruz, because no substantial gures are available. However, taking into consideration the cosmopolitan distribution of several groups, the 2000 species of micromycetes known from Mexico (see above) probably also exist in Veracruz and also represent 10% of the total; this means that there could be 20 000 species of micromycetes in Veracruz. Thus we have in Veracruz 14 000 species of macromycetes and 20 000 micromycetes, but if we reduce that number by 20% to eliminate possible synonymous species and names of anamorphs, we have 27 200 species. Extrapolating these from the 71 900 km2 area of the State of Veracruz to the 1 970 000 km2 area of the country yields a gure of 745 257 species. However, it is appropriate to divide this amount by two, because the high diversity in Veracruz is not typical for the whole country. If the number is reduced by a further 30% for repeated species, 260 840 species could occur in Mexico, according to this method of estimation. The average of these three calculations, 200 000, 100 000 and 260 840 is 186 964. I therefore propose 200 000 as the rounded gure for the number of species of fungi that grow in Mexico. This estimate is likely to be conservative because of the high biodiversity of the country. Accepting that the known species of fungi from Mexico is 7000, as discussed earlier, this means that only 3.5% are known. This gure clearly resembles that of the known species of fungi in the world, about 4.5% of the 1.5 million species estimated by

Inventorying the fungi of Mexico


Hawksworth (Hawksworth, 1991; Hawksworth and Kalin-Arroyo, 1995). It is interesting to observe that Hawksworth based his estimates on surveys of the ratio between fungi and vascular plants in well-studied areas, such as the UK where it is 6 : 1; if this ratio is applied to Mexico with their 30 000 vascular plants, we have the gure of 180 000 species of fungi. This suggests that the estimates presented here for the Mexican mycobiota are reasonable. However May (1991, 1994) suggested that Hawksworth's gure could be too high but this could reect a lack of familiarity with the behaviour of fungi in the tropics (Hyde and Hawksworth, 1997). On the other hand, Cannon (1997), in his account on the Phyllachoraceae of the world, presented the gure of 9 900 000 species of fungi in the world. Final considerations and conclusions Certain basic assumptions must be made in order to understand the diversity of fungi in Mexico discussed above: the state of our knowledge, and the methods of study. First, it is necessary to emphasize that there are as yet no intensive studies of Mexican fungi, in spite of the signicant progress made in the last 30 years. The possibility of completing and inventorying the Mexican fungi is low, because of their high diversity, the low percentage known, and the few mycologists in the country. This reality contrasts with the long tradition of several ethnic groups of knowledge of the fungi throughout the country. These people know how to recognize mushrooms, their uses, and their ecology and phenology, and even some moulds useful to humans; for example they know that some mushrooms grow in summer, but others in spring and others in winter; that Cyathus opens its cups before the rain, announcing the onset of the rainy season; and that the green mould (Penicillium) on food is good for sickness and wounds. Some of these ethnic groups say, for instance, that when collecting fungi it is necessary to be very careful in order not to damage `the roots' (i.e. mycelium), which is true; another good example of this knowledge relates to the question: What are the fungi? In an ethnic experiment (Mapes et al., 1981), a range of pictures of plants, animals and mushrooms were shown to an old Indian man and he was asked to make two groups: plants and animals. He formed three groups, those and the fungi. When asked why, and, what are the fungi?, he wisely answered `mister, fungi are fungi'. In a checklist of the common names of the fungi in Latin America (Guzman, 1997), there are more than 1000 indigenous names related to edible, medical or sacred fungi. It is essential that we collect all available information before these ethnic groups become extinct and the information becomes unavailable, as pointed out by Subramanian (1982) in India. It is known that the current state of our knowledge of the biodiversity in the world is still poor, mainly in the tropics (Heywood and Watson, 1995); it is necessary to understand the structure and functioning of ecosystems, particularly those in the tropical regions, to understand the importance of tropical diversity (Subramanian, 1982; May, 1990; Hawksworth, 1991, 1993a; Gaston and May, 1992; Harper and Hawksworth, 1994; Cannon, 1997; Hyde and Hawksworth, 1997; Korf, 1997; Rossman, 1997). It is a great challenge to raise the level of awareness of fungi, and international collaboration is required to study fungal diversity. Usually it is said that the tropics are `terra incognita', but concerning Mexico the reality is that almost all the country is `terra incognita', including the temperate regions and the coniferous forests so far explored. There are still many undescribed species, and biodiversity must be treated more seriously as a global resource. It is necessary that young mycologists take up the challenge to study the tropical fungi.



Such research will require intensive exploration to collect specimens and to survey all habitats (Subramanian, 1982; Korf, 1990, 1997; Gaston and May, 1992; Kaesuk-Yoon, 1993; Renner and Ricklefs, 1994; Hyde and Hawksworth, 1997). For this reason we need to maintain the ecological equilibrium of communities that have been lost in many regions. Fortunately the world's scientists are investigating ways to nd out how they can accelerate systematic inventories (Heywood and Watson, 1995); and conduct ambitious programmes aimed at providing a global perspective of biological diversity. Such initiatives as the Ecosystem Function of Biodiversity Programme of SCOPE and Systematics Agenda 2000 International require a high level of support (Kaesuk-Yoon, 1993; Rossman, 1994; Cannon and Hawksworth, 1995; Hawksworth, 1995a,b). Mexico, despite the contributions of CONABIO and CONACYT bureaux, established by the government to nance scientic programmes, is not able to resource intensive biodiversity inventories. Even three decades ago it was dicult or impossible to obtain funding for these kind of studies. To complete an inventory of all fungi in Mexico in a reasonable time scale would involve a major increase in systematic work. In spite of the fact that within the last ten years there have been major developments in systematic studies in the country (e.g. 389 species of ascomycetes were recorded between 1983 and 1996, as opposed to 261 from the last century to 1982 (Medel et al., 1997), we still know only a little about the fungi which occur here. Between 1985 and 1994 there were 835 additional species records or new species described, based on a review of the Mexican literature, of which the Index of Fungi catalogued 162 as new species from Mexico during 19811990 (Hawksworth, 1993a). If we have only 7000 known species of fungi after approximately 100 years, at this rate we will need more than 2000 years to discover and document the approximately 200 000 species of fungi from Mexico. An analysis of the productivity of Flora Neotropica shows how urgent it is to increase the rate of production of monographs; there are only nine monographs on fungi from the region (Prance and Campbell, 1988) and these reported only 949 species of the 50 000 they believed to be growing in the Neotropics. Surely there are more if we compare that gure with the Mexican fungi. For Flora Neotropica when extrapolated, would show that it will take more than 948 years to monograph all the fungi on the Neotropics. Nishida (1989) listed the publications of the mycologists in the Neotropics, and according to her, it seems that we have the bases, but we need to enhance, coordinate and connect all the eorts. The main problem of documenting the fungi of Mexico is to train more specialists. The present number is alarmingly low; even in the USA and Europe, systematic positions are in decline (Burdsall, 1990; Gaston and May, 1992; Hawksworth, 1993a; Hawksworth and Ritchie, 1993), except for molecular biologists discussed below. It seems that the role of the taxonomist is disappearing at a time when the practical task of identifying and recording species diversity might be better managed (Subramanian, 1982; May, 1994). But one of the problems in training specialists in fungal taxonomy is the lack of employment for them (Isely, 1972; Prance and Campbell, 1988; Burdsall, 1990; Korf, 1990; Gaston and May, 1992; Kaesuk-Yoon, 1993). Since 1960, more than 60 students in taxonomy of fungi have been trained in my laboratory, but no more than ten are now working in mycology, and those not necessarily in taxonomy. Another pressing need in the inventory of fungi is to encourage monographs of genera. In addition to nding and describing new species or new records for the country, we need to work with species complexes in order to clarify biological and ecological relationships. Rather than small regional monographs, there is a need for larger monographs, as noted by Korf (1990, 1994) with the ascomycetes.

Inventorying the fungi of Mexico


In the meantime, a realizable compromise is needed to protect the natural vegetation (Gomez-Pompa, 1987; Wilson, 1988; Halter, 1994; Gomez-Pompa and Dirzo, 1995). Land could be clearly divided into areas for agriculture and ranching purposes, and areas for conservation; these should not be mixed as they now often are. Unfortunately this view is contrary to that of some other specialists, even governments, that claim to use the areas for conservation as `little' zones for exploitation and even for grazing. It is not rational to continue opening new lands for agricultural or cattle ranging purposes, if at the same time the lands have to be abandoned after becoming exhausted through use. The problem with vegetation conservation is exacerbated by our lack of knowledge; ideally it is necessary to know which species are present in order to know how to use or conserve them. Because of the possible loss of species, the cataloguing of species before they disappear and preserving their germ plasm by fungal cultures can be justied (Korf, 1990, 1994, 1997; Guzman, 1994b; Renner and Ricklefs, 1994; Hawksworth, 1995b; Guzman et al., 1996, 1997). Environmental conservation programmes urgently need to develop at the same time as complementary training programmes for specialists. The few research institutions that have postgraduate training programmes on natural resources pay scant or no attention to systematics and are being replaced by molecular biology, genetic, biochemical and ecological studies. It is important to understand that all biological sciences depend on names of species for communication about and retrieval of all research results. That systematists concerned with species identications receive only a relatively small amount of support in many institutions, further discourages students. New methods in biology are important, but they are complementary and yet have decreased rather than increased the production of monographs because of competition from other areas of biology (Isely, 1972; Tehler, 1988; Korf, 1990, 1994; Hibbett and Vigalys, 1991; Hibbett, 1992; Hawksworth, 1993b; Petersen and Hughes, 1993; May, 1994; Mishler, 1994). It is important to note, however, that monographs based on those methods with a careful study of morphological features are needed to enhance our knowledge of all groups of organisms. But we need to be very careful in accepting conclusions based on narrow or confused taxonomic-morphological concepts, or where the identication of the taxa is not correct, even in those studies employing `stronger stringency' (Petersen, 1995). Existing institutions with specialists in systematic mycology and reference collections need to be enhanced. Collections need to be programmed for their growth and structure, and not in a haphazard manner depending on the interests of their curators. It is urgent to create networks and standard practices between collections, in order to avoid them turning into museums of themselves (Alberch, 1993; Guzman, 1994b). Moreover, systematic research centres are rare in tropical regions and very few mycologists live in the tropics (Hawksworth and Ritchie, 1993; Hawksworth et al., 1997), as is the case in Mexico. The long-term continuity of research centres poses another problem. In the last decade at least four national Mexican research institutions were closed or reduced in their research programmes, among them a tropical agricultural school and two institutes on biotic research on the tropics, with serious consequences for all the biological research programmes in the country. Although scientists have been studying fungi for many years, 95% still need description. Now with the impulse of new methods such as molecular biology, mating, biochemistry and cladistic programmes, how much longer will the task take? How far away is the end? The new methods involve relatively few species and require high levels of support and limited collections. However, monographs on genera or other systematic groups embrace



many species (many of them represented today in collections), and these studies are relatively inexpensive. It is necessary to place more emphasis on alpha taxonomy, to know rst the species of a region or a group, and then to deepen that knowledge by modern methods. Acknowledgements I am grateful to P.F. Cannon, K.L. Gaston, D.L. Hawksworth, D.S. Hibbett, R.P. Korf, R.M. May and A.Y. Rossman for providing important bibliographic references. To my colleagues V.M. Bandala, L. Guzman-Davalos, R. Medel, L. Montoya and N. Ogata, I give my thanks for critical observations. Funding was provided in part from CONABIO, CONACYT, SNI and Instituto de Ecolog a. D.L. Hawksworth, R.P. Korf, J.M. Trappe and A.L. Welden kindly critically revised drafts of this. Particularly, I wish to express my thanks to D.L. Hawksworth for his encouragement and help. References
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