/ fntemational loumal ofMedicinal Mushrooms, Vo l. 2. pp. 95-l23 (2000)

-= Genus Pleurotus (Jacq.: Fr.) P. Kumm. (Agaricomycetideae): Diversity, Taxonomic Problems, and Cultural and Traditional Medicinal Uses

Gastón Guzmán Instituto de Ecología, Apartado Postal 63, Xalapa, Veracruz .91000, Mexico

ABSTRACT: The taxonomic difficulties in the identification of the species of P le urotu ~ -are extensively dis ­ cussed. The problems are attributable to the great variation and wide distribution. as well as thb. magnified use of genetic and biochemistry smdies of strains not clearly identified. More than 1000 species of PleiVotus have been described throughout the world, in more than 25 reiated and/or confused genera. However, only approximately )O valid species are recognized in Pleurows. Modern studies based on biochemical and molecular research and inter­ breeding tests are use ful if thex are based on the macro- and microscopic morphocharacters. as we!l as on ¡he coior of the basidioma. the spore print. :ind the type of hyphal system. ali of which play an irnporcant ro le in the raxo­ nornic classification of the genus. Of the more than 71 names related to Pleurorus discussed in the present arcicle. only approximately 24 are considered as valid species in this genus. More than 20 species of Pleurotus are reponed fro m Mexico. of which only seven seem valid taxa. P. oscreatus is apparently the species studied the most, but at the same time nurnerous taxonomic problems exist in its delimication. The wild-cype is unkow n in :V!exico; more­ over. ii is the most reponed and cultivated, both for comrnercial ami research uses, together wi th P. pulmonarius. P. columbinus. and P. djamor. the latter divided into three varieties. but ali of lhem are of interbreeding among them. ;:. ' The traditional uses of the species of .f'leurorus are revised. Approxi mately l 00 common names of these mush­ • 1 { rooms are known in Mexico. Severa! sp e ~ ies of Pleu rotus are used in tradicional medicine for approximately 35 .' . disorders or diseases .

KEY WORDS: Genus Pleurows. Pleurotaceae. taxonomy, diversity. wild species. new methods. cultural and medicinal uses.

INTRODUCTION Nyssa. Ostrya, Pandanus. Picea, Pistacia. Populus. Pseudotsuga, Quercus, Salix. Tilia, Ulmus, and The species of Pleurotus (Jacq.: Fr.) P. Kumm. Wisteria; and P. septicus on Heteromeles. Perer­ [Pleurotus (Fr. ) Quél.] have been used as food or sen and Ridley ( 1996) and Petersen et al. ( 1999) fo¡ medicina.l purposes for a long time, and at reported P. djamor (or P. opuntiae) as the prob­ presem they.have an important ~ole as commercial able cause of a plant disease in New Zealand edible mushrooms. In addition, Pleurotus has forests. Species of Pleurotus, such as P. comu­ been reported as parasitic on several trees. Farr copiae, P. cystidiosus. P ostreatus, P strigosus, et al. (1989) reported the following 7 species on and P. subareolatus, together with species of 30 different hosts: P albolanatus on Pseudotsuga; Hohenbuehelia, are known to attack and consume P cystidiosus on Acer, Liquidambar, Populus, living net1Íatodes. through speci.al structures named Quercus, and Salix; P dryinus on Carya, Liq­ microdroplets, as studied by Barron (1977), Bar­ uidambar. Malus, and Quercus; P. Levis on Acer. ron and Thom (1987), Hibbett and Thom (1994), Betitla, Juglans, Liquidambar. and Salix; P. minu­ and Thom and Barron (1984). tus on Betitla, Carya, and Quercus; P. ostreatus Today high-technology commercial cultiva­ on Abies, Acacia, Acer. Alnus, Betula, Carpinus. tion of several species of Pleurotus on different Carya, Castanea, Laurocerasus. Liquidambw: agricultura! wastes and on several lignocellulosic le l Uriodendron, Lupinus, Magnolia, Malus. Morns. products has great importance in many parts ofthe J .~ i ..t 1521-9437/00/$5.00 © 1000 by Begell House. !ne. 95 '- \,, J world. This is attributable to its broad adaptabiliry (199 1) found that P. ostreatus, P. florida, P. opun­ to several substrares for the digestibiliry of ligno­ tiae, and P. sajor-caju present energy values of 1 " ; cellulosic marerials and irs potential in mushroom 265-367, whereas for Agaricus bisporns, Buswell cultivarion as well as use in animal feedstocks and Chang (1993) reponed 328-381. As P. ostrea­ .. (Chang and Hayes, 1978; Zadrazil and Kurtzman, tus is the most common species considered in the 1982; Quimio. 1986; Chang and Miles, 1989; literature, and apparently the best known, this Chang, l 991; Buswell and Chang, 1993; Guzmán name often has been used indiscriminately and it et al.. l 993a; Lelley and Janben. 1993; Royse. has been confused with other species, such as P. 1997). The oyster mushroom, i:he.English common columbinus. P. pulmonarius (frequently reported name of the Pleurorus spp .. mainly P. osrreaws. as P. ostreatus var.jl.orida), P. djamor (as Pflabel­ was first cultivated in the United States at the latus), and P. sajor-caju (a confused name). Also, beginning of the lasr century and rhen was imro­ there are severa! misidemifications of Pleurotus. duced to Europe and India (Chang, 1993: Gunde­ Kafu:ik (1992) commented that P. comucopiae, Cimerman, 1999). In Mexico the commercial P. pulmonarius, and P. salignus ha~e often been cultivation of Pleurotus ostreatus started in 1974 regarded as forms of P. ostreatus ~ltj}ough they with mycelium from Europe (Martínez-Carrera et are different species. P. comucopiae is erroneousl~ al., 199 la). World production of the oyster mush­ named "P. cornucopioides Pers." (e.g .. Zadrazil, room is rapidly increasing because the culture is 1978), while Craterellus cornucopioides (L.: Fr.) very promising in the subtropical and tropical Pers. is a mushroom completely independent of regions. From 1986 to 1994 it increased from Pleurotus. The erroneous name "Pleurorusflorida" 169,000 wns to 797,000 wns (372% increase); introduced by Eger and co-workers in 1979 (see China was the main region responsible fo r this later) is still used by many mycologists, such production (Chang,- 1991, 1993; Royse. 1997). as B uswell and Chang ( 1993 ), Peberdy et al. However, the typical º'button mushroom·· [mainly (1993), Minar et al. (1993), and others: the same A.garicüs bisporus (J. Lge ) Imbach. also known' holds for the erroneous concept of P. sajor-caju as A. brunnescens Peck] (Singer. l 986. p. -+86 ), in commercial cultures . P. sapidus is an obscure is rhe most important cultivared mushroom in the synonym for P. osrreatzts and also frequemly re­ world, fo llowed by Lentinus edades (Berk. ) Sing. ported as a valid species as discussed late r. and Vo lvariella volvacea (Bull.: Fr.) Sing. ( Chang, Anderson et al. (1973) claimed that P. sapidus has 1993). Although be tween l983 and 1984 Pleuro­ a lilac spore print, in contrast with P. ostreatus, in tus ranked sixth in world production rChang and which it is cream colored. However, the color of Miles. 1989), ir is gradually displacing the others the spore print in P. osrreanis is a confusing sub­ (Chang, 1991 ), because P'.eurorus spp. are easier ject. It seems that the P. sapidus of Anderson eta!. and cheaper to cultivare; at present, ir is third in ( 1973) is P. ostreatus. and their P. ostreatus is P. world production (Chang et al. , 1993a: Peberdy pulmonarius, as observed by Bresinsky et al. et al., 1993). Moreover, several research studies ( l 987). According to Singer (personal communi­ have focused on t4e culture óf Pleurotus (e.g., cation, 1975), Phillips (1981 ), and Wakefield Eger et al., 1974~ Zadrazil, 1974; I-:Channa and and Dennis ( 1981 ) the color of the spore print of P. Garcha, 198la.b; Volland-Nail, 1981: Hilber, ostreatus is lilac, but Bresinsky et al. ( 1976) con­ 1982; Zadrazil and Kunzman. 1982; Kunzman and sider it to be whitish to cream with slight incarnate Zadrazil, 1982; Bresinsky et al., 1987; Laborde, tinges. Moreno et al. ( 1986) reported the spore l 989; Chang, 199 l; Petersen and Hughes. 1993 ; print of P. ostreatus.as whitish to yellowish-cream. Vilgalys et al .. 1993, 1996; Perersen and Gordon. Hilber (1982) has shown that collections with 1994; Vilgalys and Sun, 1994a.b: Labarere and lilaceous and white spore prints are conspecific Irac,:abal, l 995; Petersen, 1995a: Gibriel et al.. among them, a position supported by Watling 1996: Petersen and Ridley, 1996: Müller, 1997; and Gregory (1989). Smith (1978) determined Scherba et al., l 999). that ·the spore print of P. osrreaius is whitish at With regard to the edible properties of Pleuro­ fi.rst, then changes to lilac gray after drying. Bas tus, Crisan and Sands (1978), Bano et al. (198 l ), ( 1990) considered the spore print of P. osrrearus as ., Bano and Rajarathnam (1982), and El-Kattan et al. whitish. pale gray, pale lilaceous gray or pale

96 .,i

,. olive-gray-buff. Recently Perersen and Krisai­ related with other genera such as lentinus and ... Greilhuber ( 19Q6) considered the spore print of Hohenbuehelia. In modem books, for example, the neotype of P. ostreatus grayish. Hawksworth et al. ( 1995), Omphalotus olearius All the cultured basidiomara of P. ostreatus (DC.: Fr.) Sing., a well-known agaric belonging to fromEurope obtained by the author's co-workers Paxillaceae (Singer, 1986), is still considered (Guzmán et aL 1993a, 1994) presemed a lilac Pleurotus, following the old concept of Gillet in to lilaceous spore print, even rhose identified as 1884 (!) (Heim, 1957). The genus Hohenbuehelia P columbinus. whereas those of P pulmonarius is well separared from Pleurotus by its rhick­ presented a white spore print. Severa! commer­ walled cystidia known as meruloids. Nonetheless. cial cultures of Pleurotus reported as P. ostreatus Singer (1986) and Pegler (l 983a) classified as acrually belong ro other species. P columbinus, Pleurotus species with metuloids, for example, P. pulmonarius, "P. sajor-caju." "P. ostreatus var. P. j loridanus and P. rubarius. As for Lentinus, ac­ florida," "Pflabellarus," P. djamor, and others are cording ro Pegler (1977 a,, l 983a,b, 1986), it is 1 confused with the cultures of P. ostreatus. Also, separated from Pleurows by r~e strucrure of the the author and his co-workers found that a strain hyphal system, dimitic in Lentinus and monomitic of Pleurotus sp. sent to Mexico from Argentina in Pleurotus, resulting in a texture that is tóugh in produces basidiomara belonging, to Schizophyllum Lenrinus and fieshy in PleLLrotus. However, there commune Fr.: Fr ( !). are rrue species of PleLLrotLLs with a dimitic system All the rnisidentifications and problems of as discussed later. Pleurotus are attriburabie mainly to the lack of Genera related ro Pleurotus, with the excep­ taxonornic studies on the genus, and rhen to the tion of the anamorphic states (e.g. , Antromy­ difficulty in identifying the species and the fre ­ copsis Pat. et Trnb. and Pachyma Fr.) are listed quent use of interbreeding tests ro define species. in Table l (Konrad and Maublanc (1924-1937); Guzmán et al. ( 1994) found rhat the reports of P. Pilát, 1935; Singer. 1949. 1962. 1975, 1986; ostreatus from Mexico. such as those of Guzmán Heim. l 95Tl. Hohenbuehelia is accepted by rhe ( 1977 a), Martínez-Carrera and Morales (1988), majority of authors as an independent genus in and Martínez-Carrera et al. (1988a.b), in actuality Agaricales (e.g .. Thom, 1986). Nothopanus was refer ro P. djamor. Considering rhe grear tradi­ considered '.lil independent genus by S inger (1949, tional and commercial importance of the species 1962. 1975), but Horak (1968) and Singer (1 986) of Pleurorus, the present article considers how to stated that this is a synonym of Pleiirorus fol­ understand the taxonornic problems related to the lowing Dennis ( 1953, 1970). Petersen and Krisai­ genus, the detlnition of rhe genus, the best way to Greilhuber ( 1999) and Petersen et al. ( 1999) identify rhe species, and the revision of the Mex­ designated Nothopanus (Singer, 1944, type N. eLL­ ican species and their uses and culture. grammus) as a synonym cif Pleurotus but they described the genus Neonothopanus. with the type N. nambi (= Pleurotus nambi) to avoid confusion MATERIALS· ANO METHODS with the Nothopanus of Singer that is another mushroom for Petersen and Krisai-Greilhuber. An.extensive critica! revision of the bibliog­ There are two species in Nothopanus sensu Singer raphy on Pleurotus (more than 230 references) is common in the neotropic: N eugrammus and N hy­ the basis of this article. as well as the experiences grophanus. Pegler (1983a) consideredNothopanus of the author over severa! years of studying their from the Ca,ribbean region. Guzmán (1977a, 1983, taxonomy, culture. and traditional uses of rhe 1997) discussed both as Pleurotus from tropical species of Pleurotus. Mexico. Since the lamellae are very distant and sometimes forked or interveined, and the basid- ,ioma irregularly mottled of vinaceous red color, it PROBLEMS IN THE GENUS PLE.UROTUS is better to consider these mushrooms as independ­ ent genera of Pleurotus. Petersen and Gordon Consensus regarding generic lirnits of Pleu­ (1994) described P hygrophanus as Pleurotopsis -:. · rotus has not been reached. and ir is frequemly guatemalensis from Guatemala (also reponed

97 TABLE 1 of Pegler and others. Pegler and Young (1 983) Sorne Genera Related to Pfeurotus (Many of · separare Pleurotus from Lenrinus by the lengrh of them now belong to another genera or are rhe skeletal hyphae, up to 300 µm in Pleurotus. independent; see text. The anamorphs are not Romagnesi ( 1969) was one of the first to observe considered.) rhe importance of rhe presence of the sclerified Acanthocystis (Fay.) Kühn . hyphae in Pleurorus: however. Bresinsky et al. C!itocybe (Fr. ) Staude ( 1976) and others considered such a criterion as Crepidotus (Fr.) Staude only of limited value. because rhey assumed rhat Faerberia Pouzar the developmem of sclerified hyphae depended on Geopetalum Pat. the type of substrate on which rhe species grow. Hohenbuehelia Schulz. Another problem in Pfeurorus is its raxonomic po­ Hypsizygus Sing. Lampteromyces Sing. sition. Konrad and Maublanc ( 1924-1937), Singer Lentinus Fr.' ( l943, 1949), Heim (1 957), and Dennis (1970) Lentodiopsis Bubák considered Pleurotus in Agaricales, Tricholomat­ Lentodiellum Murr. aceae. Notwithstanding, on rhe b a~is of rhe hy­ Neonothopanus Petersen et Krisai-Greilhuber phal system, Singer (1962, 1975. 1986) and Peg-ler Nothopanus Sing. Omphalotus Fay. ( l 977a) considered Pleurorus belonging to rhe Ompha/opsis (Noordel.) P.D. Orton fa mily Polyporaceae bur in Agaricales. Kühner Ossicaulis Redhead et Ginns and Romagnesi (1953) considered Pleurows in the Pane/lus P. Karst. Pleuroracées in ¡he Agaricales. Jülich (198 1) ac­ Panus Fr. cepted fo r Pleurotus rhe family Pleuroraceae fol­ Phylloporus Quél. lowing in part Kühner and Romagnesi (1 953 ), Phyllotopsis J.-E. Gilbert et Oonk ex Sing. Pleurocol/ybia Sing. but in rhe Polyporales. Watling and Gregory ( 1989) Pleurotellus Fay. considered the farnily Pleuroraceae in the agaric Pleurotopsis (Henn.) Earle ílora of Great Brirain, :.is did Imazeki ;ind Hongo F?hodotus R. Maire ( 1983) in Japan. Pegler ( l 983a) placed Pfeu rorus Tec tella Earle in the Aphyllophorales, Polyporaceae. and larer (Pegler, 1986) changed rhe family for Lentinaceae. The posiüon most widely followed by specialists later in Petersen, 1995b). However, Petersen (per­ is that of Singer (1962: 1975. 1986) to place sonal communi_9tion. 1997) maintained that P. Ple uro rus in Agaricales, Polyporaceae (e.g .. Moser. guatemalensis is~ really Plicatura obligua or a 1983; Kaarik. 1992) or thar of Pegler (1 977a) to similar species. The genus Plicarura was de­ place Pleurotus in Poriales, Polyporaceae. scribed by Peck (1872) based on P. alni Peck, a The hyphal system. either mono mi tic or dimi­ mushroom similar to Cantharellus on the basis of tic; cy lindrical spores and hyaline. nonglobose the folds in the hymenium. For Hawksworth et al. generative hyphae: pleurotoid or t1abellare habit: (1995) this genus belongs to the Corticiaceous not well developed stipe; and white, whitish, gray­ mushrooms. Singer (1986) excluded this genus is h, grayish blue, lilac, or pink spore print are the from Agaricales. Concerning Lampteromyces, it main features defining Pleurotus. Regarding rhe was based on Pleurotus japonicus, studied by division of the genus following Singer (1986), Singer (1943). P. japonicus is a poisonous mush­ there is still confusion. Singer (1986) divided the room that Singer (1986) considered belonging to genus into the sections: Lepiotarii (Fr.) Pi!. , Ca­ Lampreromyces. lyprrati Sing., 'Pleurow s, Coremiop!eurotus (0 . One of rhe first taxonomic problems regard­ Hilber) S ing., Lentodiellum (Murr.) S ing., and ing Pleurotus is its correct delimitation and taxo­ Tuberregium Sing. It is very difficult ro follow this nomic position. There is no agreement among divisi on, however, because of inconsistent fea­ the specialists, because for Singer (1949, 1962, tures and/or their repetition, for example, Lepio­ 1975, 1986) Pleurotus presents a broad concept, tarii and Calyprrati are disringuished by a veil. but with both a monornitic and a dimitic hyphal sys­ it is also present in P. sajor-caju, considered in tem, in contrast to the previously discussed views Sect. Lentodiellum where there are species with-

98 our an ~mnulus, for example. P levis and P hirtus. others, this is assigned to the Hyphomycete genus Hi!ber (1982, 1989, 1997) and Zervak.is ( 1998) A.nrromycopsis: A. macrocarpa (A. broussonetiae) considered rhe Seer. Coremiopleurotus as a sub­ in the first and A. guzmanii (A . smithii) in the genus of the well-developed synnematous anamor­ second (Guzmán et al., 1980, 1991; Moore, 1984; phic stare, as is rhe case fo r P. cystidiosus and P. Stalpers et al., 199 l; Zervak.is, 1998; Capelari, smúhii, shown by Pollack and Miller (1976) and 1999) (see Table 2). It is interesting to observe Guzmán er al. (1980, 199 1), respecrively. In that in sorne cultures of P. osrreatus with a high Pleurotus cysridiosus rhe anamorph state is based , concentration of C02 sorne abnormal basidio­ on the asexual forms observed by Kaufert ( 1936) mata with defo rrnations similar to Antromycopsis and Semerdzieva (1965) in P. corticatus s. auct, sinnema are frequently obtained, such as those non s. S inger ( 1986) (see later). Zadrazil and observed by the author in the cultures of his Kurtzman ( 1982) observed rhe problem of placing co-workers (Guzmán et al.. 1993a) and by Zadra­ P cysridiosus in one of the sections of Singer's zil (1974, Figs. 11-1 and, 12), Zadrazil (1978, (1986) classification. Pilár (1935) divided the Figs. 16--4 and 17 ), md Za~ra zi l and Kurtzman subgenus Eupleurorus Pil. imo 5 sections, as (1982, Fig. Sf 4). Also. Desbiens et al. (J 978) ob­ Acanthocystis, Omphaliopsis, Paneilus. Phyllo­ served that a Pleurows sp. they cultured were sen­ to , wpsis. and Rhodorus now belonging to anorher sitive conc emrations of C02 and developed ab­ genera. Konrad :md :'vlaublanc ( 1924--1 937) sub­ norrnal fruiting bodies. both in macro- and divided Pleurorus imo tive subgenera. Hilber microscopic fearures. Little is known about why ( 1982. l 997) divided P!eurorus imo three sub gen­ or when Pleurorus forms its asexual state. The era: Pleurows. Coreomiopleuroms O. Hilber, and aurhor had observed the A.mromycopsis srate of Lentodiopsis (Bubák) O. Hilber. P. smithii in culture and in wiid conditions. The Regarding rhe anamorph state of Pleurows. variarion in the form and color of the basidio­ ir is known that in P L'ysridiosus. P. smithii. and mata under culture led ro a consideration of such

TABLE 2 Species ar Names of Pleurotus and Anamorphic States Discussed in this Article (Those in bold letters are valid ar possible valid taxa.)

Antromycop$iS broussonetiae Pat. et Trab. (A macrocarpus) A. guzmanír_~alpers, Seifert et Samson (A smithít) (anamorph of P smíthit) A. macrocarpa Stalpers. Seifert et Sampson (A. broussonetiae) (anamorph of P cystidiosus) A. smithii (Guzmán) Guzmán et R. Valenz. (A guzmanit) P. abíetico/a Petersen et Hughes P agaves Dennis P. aba/onus Han , Chen et Cheng (a species related to P cystiodíosus and P smithit) [according to Hilber, 1997, is P cystidios1,1s subsp. aba/onus (Han, Chen et Cheng) O. Hilber] P albellus (P.at.) Pegler, seems clpse to P elongatipes P albolanatus (Peck) Kaufm., not well known species P. bajocalífornicus Esteve-Rav., G. Moreno et N. Ayala P. calyptratus (Lindbl. in Fr.) Sacc. (Sokól and Szczepka, 1995) P calvescens (Berk.) Sing. (Lentinus striatulus Lév.) (see P focket) P calyx (Speg.) Sing. (Lentínus patulus Lév.) P catephes (Berk.) Sacc., a poorly known species P. columbínus Bres. (P ostreatus var. columbinus about severa! authors) P. cítrínopíleatus Sing. (P cornucopiae var. citrinopíleatus) P concavus (Berk. ) Sing. [Lentinus concavus (Berk.) Henn.; L. concavus (Berk.) Comer; Lentodiellum concavum (Berk.) Murr.] P. cornucopíae (Paul. : Pers.) Roll. P cornucopiae var. citrinopileatus (Sing.) Ohira (P citrinopileatus) P corticatus (Fr.: Fr. ) Quél. s. str., conspecific with P dryinus P corticatus sensu Kaufer 1936 is P cystidiosus Continued

99 TABLE 2 (continued)

P. cystidiosus O. K. Mili. (P corticatus ss. Kaufert) (anamorph: Antromycopsis macrocarpus) P. cystidiosus var. formosensis Moncalvo P. djamor (Fr.) Boedijn var. djamor (P flabellatus, P mexicanus) P djamorvar. roseus Comer (P djamorvar. djamor) P djamorvar: salmoneostramineus (L. Vass.) Guzmán (P sa/moneostramineus, P djamorvar. djamor) P. dryinus (Pers.: Fr.) P. Kumm. P dubius (Lloyd) O.A. Reid (Stereum dubiumlloyd; Hohenbuehelia?), a not well known species P elongatipes Peck [Hypsizygus tessullatus (Buil.: Fr.) Sing.; see P. albellus] P. eryngii (OC.: Fr.) Ouél. P eugrammus (Mont.) Oennis [Nothopanus eugrammus (Mont.) Singer; Lentinus eugrammus Mont.] P. eous (Berk.) Sacc., a synonym of P. djamor P. euosmus (Berk.) Sacc. (confused, related to P ostreatus or P. cotumbinus) P feru/ae Lanzi (see Appendix) P. flabellatus (Berk. et Br. ) Sacc. (P. djamorvar. djamor) P. f/avotanatus (Berk. et M.A. Curt.) Sacc., poor known species P. floridanus Sing. (non florida s. au et.) P. florida Eger nom. nud. (P. ostreatus var. florida Eger; P. ostreatus or P. putmonarius) P. fockei (Miq.) Sing., a not well known species P. tuscosqua mulosus Reid (clase to P. abalonus, P. cystidiosum, and P. smithii) P. hirtus (Fr. ) Sing. [Lentinus scteropus (Pers.) Fr.] P. hygrophanus (Moni.) Oennis [Nothopanus hygrophanus (Mont.) Sing.; Pleurotopsis guatemalensis Petersen et Gordon] P. japonicus Kawam. [Lampteromyces japonicum (Kawam.) Sing.] P. komarnitzkyiVassilk., with an uncertain taxonomic position (Hilber, 1982: Singer, 1986) , however, recognized by Hilber (1997)

P. laciniatocrenatus (Speg.) Speg. 1 P. /evis (Berk. et M.A. Curtis) Sing.: Lentinus /evis (Berk. et M.A. Curtís) Murr. P. lutealbus Beeli P. mexicanus Guzmán (P. djamor var. djamor) P. mmutus Peck (Crepidotus minutus (Peck) Murr.] P nambi (Speg.) Sacc. [Nonothopanus nambi (Speg.) Petersen et Krisai-Greilhuber] P o!earius (OC.: Fr. ) Gill. [Omphalothus olearius (OC.: Fr.) Sing.] P. opuntiae (Our. et Lév.) Sacc. (P yuccae) P. ostreatus (Jacq.: Fr. ) P. Kumm. var. ostreatus "P ostreatus var. apr}atachiensis" Hilber. 1997 P. ostreatus var. florida Eger (P. florida; P ostreatus or P pulmonarius) P ostreatus var. columbinus (Quél.) Qué!. (P columbinus) P ostreatoroseus Sing. (P. djamorvar. djamor, P. r oseopi!eatus) P. populinus O. Hilber et O.K. Miller (belongs to the P. ostreatus complex) P. pulmonarius (Quél.) Sing. (P. ostreatus var. florida?, P sapidus) P. roseopi!eatus Sing. nom. nud. (P djamorvar. djamor) P. sa/ignus Fr. , a not well known species P. salmoneostramineus L. Vass. (P. djamo'r var. sa/moneostramineus) P sajor-caju (Fr.) Sing. (Lentinus sajor-caju) (not P sajor-caju s. auct.) P. sajor-caju s. auct. (P. ostreatus and P. pulmonarius) P sapidus (Schulz. apud Kalchbr.) Sacc. (? P. cornucopiae;? P ostreatus;? P pu/monarius) P. septicus Fr.: Fr. (uncertain species) P. serotinus (Schrad.: Fr.) Pat. (Panellus serotinus Pers. sensu Sing., sensu Kühn.; Crepidotus, Murr.; Sarcomyxa, P. Karst.) P spodoleucus (Fr. ) Quél., a not well known species (? P. ostreatus) P. smithii Guzmán (anamorph: Antromycopsis guzmani1) P strigosus (Berk. et M.A. Curtis) Sing. (a synonymous of P levis) P subareo!atus Peck, a poorly known species (? P. pulmonarius) · P. tubarius (Pat.) Pegler P. tuber-regium (Fr. ) Sing. [Lentinus tuber-regium (Fr.) Fr.; Panus tuber-regium (Fr.) Comer] P. u/marius (Bull .: Fr.) Quél. P. viscidus Harmaja, non P viscidulus (Berk. et Br.) Clel. from Australia, that seems a Hohenbuehelia. P. yuccae R. Maire (P. opuntiae)

100 ·-= changes as indicative of another or new taxa. Bre­ ered S species from Great Britain: however, Rea sinsky et al. 0976, 1987) observed that P. ostrea­ ( 1922) reponed 43 species, most of them now be­ tus, P columbinus. and P pulmonarius presented longing to other genera (this example shows how large variations of the basidiomata under culture. the magnirude of the genus can change). Zervakis Another anamorph genus reported in Pleurotus and Balis ( 1991 ). on the basis of morphological is Pachyma. in Pleurotus tuber-regium (Singer, characteristics, including the hyphal system, rec­ 1986). However. Pachyma is also reported as the ognized 4 species in Greece, but later ( 1996) recog­ anamorph of Paria cocos (Schwein.) Wolf, as nized 8 species in Europe based on interbreeding, Pachyma cocos (Schwein.) Fr. (Weber, 1929), although they observed that there is controversy re­ and on the other hand, the position of Pleurorus garding P. ostreatus, P. coiumbinus. P. puimonar­ tuber-regium is in discussion. ius, P. sapidus, P. comucopiae, and others. Hilber Conceming the problem of the production of ( 1978, 1982), also based on the hyphal system spores in the basidiomata of Pleurotus sp. in com­ and the development of a.sexual states, recognized mercial culture houses, Leal-Lara (1978) de­ P. ostreatus, P. pulmonarius:,p columbinus. P. cor­ scribed the chance discovery by Eger in 1970 (in nucopiae, P. opumiae, P. kom"amirzky. and P. eryn­ Arch. Mikrobiol. 74) of a strain with sporeless gii, but later (Hilber, 1997) recognized P. cys­ production in the basidiomara of P ostrearus. ridiosus with two varieties. P. ostreaw s "var. which perhaps is caused by an infecrious agent. In appalachiensis." P. popuiinus. P. dryinus, P. calyp­ comrast. Imbemon and Labarere ( 1989) and Im­ tratus. P. i!ous. P. conmcopiae var. comucopiae. bemon and Houdeau ( 1991) produced at INRA at and P. cornucopiae var. cirrininopileatus. Comer Bordeaux. France mutant strains of P. ostreatus (1 98 l ) from Malaysia described approximately 40 and P. puimonarius without or with poorly spored rnxa in Pleurotus but later (Comer. 1994) excluded production. These strains were produced by ultra­ severa! ''anomalous species," because of the ab­

violet muragenesis. Ohira ( 1979) 1 described a sence of clamp connections in them. He considered sporulation-deficient mutant in P. pulmonarius, these latter as belonging to Pleurocollybia. Pegler and Chang et al. ( 1985) and Change and Miles (1969. 1972, l977a.b. l983a, 1986) described lO ( 1989) a sporeless strain in "P. f lorida." species of Pleurorus from Africa. Antilles, India. Nepal. Pakistan. and Sri Lanka. Dennis (1 953. 1970) in Venezuela and adjacent countries reported THE KNOWN SPECIES OF 14 species of Pleura tus. sorne of them belonging m PLEUROTUS ANO PROBLEMS Hohenbuehelia. Batista-Pereira ( 1988) considered IN THE!R-CLASSIF!CATION 8 species from Rio Grande do SuL BraziL of the 21 reponed there. He considered P. osrreatus, but not The few taxonornic studies on Pleurotus are P. djamor in any of its forms. There is probably incompiete, except that of Hilber ( 1982), but deal sorne confusion, because he described a spore print mainly with European species, of which he con­ "branca a creme até cinza," which might involve . sidered apprnximately 10. Singer (1986) reported the two species. Murrill (1916) discussed 9 species approximately 40 species [bur.Singer (1943), con­ for North America (Mexico was not considered) sidered only 12 species, then (1949) 15 species, but all of them as Crepidorus. and among them one of them as Pleurorus sp., and ali of them in a C. ostreatus. C. comucopiae. and C. serotinus. dichotomic key]. Pilát (1935) described more than He also discussed 8 doubtful species, such as 60 species, but many of them now belong to other Ple uro tus "Salignus. Imazeki and Hongo ( 1983) genera. Konrad and Maublanc (1924-1937) con­ and Imazeki et al. (1988) considered only four sidered 10 species and 64 as doubtful and/or species: P. cornucopiae var. citrinopileatus, excluded species. Kühner and Romagnesi (1953) P. cystidiosus. P. ostreatus (=P. pulmonarius), and discussed 12 European taxa, while Moser (1983) P. salmoneostramineus from Japan. Cleland refered to only 8 and Bresinsky et al. (1976) 6. (1934-1935) reported 7 species from Australia. all Kafuik (1992) considered 7 species in the Nordic of them Australian species, except his "P. osrrea­ countries. Bas ( 1990) described S species from The tus." Hilber, in (1989) revising the mushrooms Netherlands. Watling and Gregory ( 1989) consid- considered as Pleurorus, found that 17 belong to

101 other genera independent of Pleurotaceae, such as The main species in the genus frequently P. tuber-regium that ac;:cording to Pegler (1983b) is discussed in the bibliography or cultured in the Lentinus. Recently, Petersen et al. (1999) consid­ laboratory or in cornmercial factories, other than ered 15 species of Pleurotus, 8 of which require P. ostreatus, are P. abalonus, P. citrinopileatus, further study. P. columbinus, P. comucopiae, P. cystidiosus, P. dja­ Accordíng to Hawksworth et al. (1995) there mor (as P. ¡1abellatus), P. dryinus, P. eryngii, P. levis, are approximately 50 valid species in the genus P. opuntiae, P. pulmonarius (as "P. florida"), and Pleurorus; nevertheless, more than 1000 narnes "P. sajor-caju," among others. P. ostreatus seems have been proposed in the genus· since more than the most studied, but at the same time the most two centuries ago (e.g ., Saccardo, 1882-1931 ). difficult to classify because of its broad distri­ However, of the 71 species discussed in the pres­ bution and variation. As example, Singer ( 1956) ent article (Table 2), approximately 24 are valid. stated that the P. ostreatus complex at the tropics Then, considering that no complete modem re­ presents a white and thin pileus and pale drab or vision of the genus Pleurotus exists, there is almost white spore prims. which' ayee with the considerable confusion on the status of several description of P. djamor (Guzmán et 'hl., l993b)! species. Singer's great book (1986) on Agaricales, Even when the author showed to Singer the type considered one of the most complete on the sub­ of P. smithii in a fresh basidioma growing in its ject. did not consider P. djamor but P. j labellatus. habitar at Mexico City, in 1974 (Fig. 1), Singer a well-known synonym of that species, was con­ stated he was sure it was P. ostreaws. however P. sidered in the same section as P. ostreallls with a smithii presents a scaly pileus never observed in monomitic system, and P. ostreatoroseus (another P. ostreatus, in addition to other microscopic dif­ synonym of P. djamor) is in a section with a ferences and the color of the spore print. Smith dimitic system. These examples show how con­ (1978) concluded that in P ostreatus there are fusing are the taxonomy and nomenclarure qf the many variants. "or closely related taxa?" He genus Pleurotus at present. recorded a whitish variant of P. ostreatus in the

2 study of the basidioma fco lor. tex ture. spore print color r morphology. macro & microscopicaly

!7 \ molecular studies 3 study of the wild substratum

TO KNOW THE SPECIES l 6 4 biochemical studies ~ isolation of a strain, obtention of basidiomata and mating with other we!l 5 identified strain ~ obtention ofbasidiomata and srudy as in l and 2

FIGURE 1. The most convenient method to fully understand the species of a Pfeurotus (in the figure, the type of P smithii in its habitat).

102 United States. However. the author-on a field trip was recently discussed by Petersen and Krisai­ in Michigan with Dr. Smith in the surnrner of Greilhuber ( 1999) in selection of the epitype from 1965-found [his whire forrn rhat corresponded Austria. The species has a depressed to infundi­ well with P pulmonarius. buliform, cream-ocher to grayish ocher pileus, eccentric stipe, pinkish violet spore print, and a monomitic hyphal system. Blaich (1976) distin­ DISCUSSIONS ON SOME SPEClES guished P. comucopiae using elecrrophoretogram analysis of enzymes. Hilber's (1982) strains of For P. abalonus see. the later discussion of P comucopiae sometimes produces an evanescent P cystidiosus. P. columbinus is frequemly consid­ veil, as noted by Petersen and Krisai-Greilhuber ered in the literature as a variety of P ostreatus (1999) in P levis under culture. Also Soba! et al. (Pilát, 1935; Eger et al.. 1979; Hilber, 1982, 1989, (1997) observed a veil in the basidiomata of Len­ 1997; Watling and Gregory, 1989; Buchanan, 1993; tinus levis developed in, the laboratory. Pleurotus Petersen et al.. 1999) or a synonyrn of P ostreatus cornucopiae sensu Guzmán (Guzmán, 1977a) (Bas, 1990). Kühner and Romagnesi (1953) and seems to be long to another species, as discussed Moser (1983) considered it as an independent in the following. species, based on the blue pileus. For Guzmán et P. calyptraws is another European species. al. (1994) it is a valid speciés. based in the nega­ with a well developed veil and dimitic hyphal tive interbreeding between strains of P ostreatus system. Recently Sokól and Szczepka ( 1995) de­ and P. columbinus. al! of them frorn Europe (one scribed this species. Singer ( 1986) considered P of P ostrearus from the U nited S tates); these strains calyptratus the only representative of the sub­ were idemified through the basidiomata obtained genus Calyptrati. The species was considered by frorn rhem. P. columbinus is defined by its bluish Singer ( 1943) in Tectella. Petersen et al. (1999) deep or blue-gray tones in the pil~us. colors not discussed that P. calyptrams presents nemarosta­ found in P. ostreatus. P. catephes is :i poorly known tic microdroplets, such as those observed in other species, not considered by severa! authors. for species of Pleurotus. ex.ample, Singer(l949. 1962. 1975, 1986), Pegler P. cystidiosus was described from the U mied (1983a,b. 1986), ::md Buchanan (1993). It was re­ States (Miller. 1969), and then reported from Africa ported by Dennis (1953. 1970) from Trinidad (Morocco) in its anamorph state (Pollack and Mil­ (Caribbean region) and Guzmán (1983) frorn Mex­ ler, 1976). P. corricatus sensu Kaufert (l936) is a ico (Penín.sula of Yucatan). For Pegler ( 1977 a) it synonym of Miller's species. A good photograph is close to"P.~opuntiae. but differs in the structure of P. corticatus sensu Kaufert by Pilát ( 1930) rep­ of the distant lamellae. resents a probable P. cystidiosus. although P. cor­ P. citrinopileatus is known only from eastern ticatus s. str. seems conspecific with P. dryinus Asia (Singer, 1943; Ohira: 1990). It is a very pop­ according to Singer ( 1986). At present, P cysti­ ular mushroom among the commercial cultures diosus is known in the southeastem United States. (Stamets, 1993). Petersen and Krisai-Greilhuber Europe. Africa, India. Taiwan, Indonesia, Java, (1999) and Petersen et al. (1999) considered this and Brazil (Han et al., 1974; Jong and Peng, species a variant of the European species P. cor­ 1975: Moore, 1985: Stalpers etal., 1991; Zervakis nucopiae, based on Ohira ( 1990) who showed fer­ et al., 1992: Moncalvo. 1995; Capelari, 1999). tility between strains of both species, in spite of Capelari ( 1999) reponed P. cystidiosus from Brazil rhe fact that the basidiorna of both are different in in both its anarnorphic state,Antromycopsis macro­ color. yellow in P. citrinopileatus, whitish in the carpa. ánd the basidiorna, which was not pre­ latter. Ohira ( 1990) named the species P. cornu­ served. Moncalvo ( 1995) described a new variety copiae var. citrinopileatus. Parmasto ( 1987), who of the teleomorph: P. cystidiosus var.formosensis. considered P. citrinopileatus an excellent edible close to P smithii according ro hirn, based on rnushroom. discussed the differentiation with differences found in the basidioma, color of the P. comucopiae on the basis of smell. which is like spore print, and in DNA srudies. It is imeresting fish in the first and anise in the latter, as well as to observe that this Asiatic rnushroom is closer to the different sizes of the spores. P. comucopiae the Mexican and South American species than

103 to the North American and pantropical species Hilber ( 1997) named this mushroom P eous with P. cystidiosus. P. smirhii, which differs from P. cys­ P. flabellatus, P salmoneostramineus, and P. os­ tidiosus in the pleurocystidia. is known from treatoroseus as synonyrns, but he did not consider Mexico and Peru (Guzmán, 1975; Guzmán et al., the name P djamor (!). Pegler (1976) considered 1980), Cuba (Rodríguez and Camino, 1990) and P. eous from India, with pink basidioma. P eus­ Argentina ( S pinedi, 1995), in both perfect and mus is another species, of doubtful taxonomic anamorph ?tates in all the repons. P. smirhii was position, reiared to P. ostreatus (Singer, 1962) or not considered by Petersen et al. (1999) . Hilber to P columbinus (Hilber. 1989). P. djamor is the (1997) considered P. smithii (wíth its anamorph valid name for P. eous, although often confused state) as a synonym of P. cysridiosus subsp. cys­ with P ostreatus, as the "tropical form of P os­ tidiosus, but he did not consider Mexico or South trearus" of S inger (1956). All the nati ve strains America in the distribution of chis mushroom reported by Martínez-Carrera (e.g., Martínez• where P. smithii is common. P. abalonus is a Carrera and Morales, l 988) as f ostreatus are mushroom from Japan and Taiwan. closely related P. djamor. E ven the pink form consitlered by Cor­ to P. cystidiosus and P smithii. It differs in the ner (1981 ) as P. djamor var. roseus ~ P djamor size of spores, up to 14 µm. long vs. up to 18-20 var. djamor because of the inrerbreeding among or 17-18 um in P cvsridiosus and P smithii, them (Guzmán et al., l993b). Also, P ostreato­ ' . ' respectively, and in addition ir does not have roseus considered by Bononi et al. ( 1991 ) and pleurocystidia (Han et al., 1974: Neda and Foru­ Cedano et al. (1 993) from Brazil and Mexico. kawa. 1987). However. Singer and Hilber ( 1977) respectively, is P djamor var. djamor (Guzmán et considered P abalonus · related to P cystidiosus. al., l993b ). P. djamor is a panrropical mushroom Far Imazeki and Hongo (l 983 ), P abalonus is a (Pegler, 1977a.b. 1983a, 1986: Nicholl, 1996). synonym of P. cystidiosus. but Imazeki et al. but it also reaches New Zealand (Petersen and ( 1988) did not consider both species. Zenvakis Hughes, 1999), where Petersen (1995a,b) and Pe­ ( 1998) and Zervakis et al. ( 1994) found that P aba­ tersen :md Ridley ( 1996) reported ir as P opun­ lonus and P cystidiosus a.re not compatible. Re­ tiae. However, Nicholl ( 1996) found that the New cently, Reíd et al. ( 1998) described P. f uscosqua­ Zealand strains are compatible with those Mexi­ mulosus from South Africa. which ir is very clase can strains identified as P. djamor. The reports of to P. cystidiosum, P. smithii, and P. abalonus. Reíd P. flabellarus by Pegler (1 977a,b, l983a) from and co-workers discussed that the South African India. Africa, and Antilles, respecrively, be long to species differs fro!!!_ those species mainly in the P djamor. according to Pegler (1986). Nicholl cheilocystidia and also in the color of the basid­ (1996) and Pe tersen (1 995a) di vided P. djamor ioma in P. abalonus, but they presented an inter­ into three variants according to the color of the esting interbreeding test among these species. pileus: (1 ) whitish, tan to pale, (2) grayish. and P. fi1scosquamulosus is not compatible with P. cys­ (3) pink. but all inrercompatible among themselves. tidiosus and P. smithii, using strains from the Indeed, P djamor presents a great variation in the Miller collection and the American Type Culture color of the basidioma, as observed by Guzmán et Collection (ATCC), respecúvely. However, it al. ( 1993b ), depending on the light of the habitar: presented 100% of compatibility with a strain of the pink forms are from the forests and the whitish P. cystidiosus from the ATCC (!). The Miller strains forms are from open areas. contrary to the opin­ of P. cysridiosus and P smithii were 17 .5% com­ ion of Petersen and Hughes (1999) and Nicholl patible, anda strain of P cystidiosus from Taiwan (1996), who stated that substrates or edaphic fac­ was 5% compatible with P. fuscosquamulosus. tors may maintáin the natural variation. Concern­ These observations of Reid and co-workers show ing the distribution of P. djamor, Vilgalys and Sun how important it is to take care in conclusions ( l 994b) found spores of this tropical mushroom in based only on imerbreeding tests. Petersen et al. Switzerland. China, Hawaii, and Canada by a spore (1999) did not consider P. fuscosquamulosus. trapping method in selected strains of Pleurotus. P djamor is the most important species in the To find spores of P. djamor in Hawaii or China tropics and subtropics, and as discussed by Guz­ seems no surprise, ·because the first place is trop­ mán et al. (1993b), was known as P. jlabellatus. ical and the second has a broad tropical and sub-

104 tropical zone in the south. But to find spores of which it differs in rhe large pileus and short stipe. P. djamor in Switzerland and Canada is surpris­ Patouillard, according to Pegler (1983b), reported ing, and perhaps an error. Probably the finding in P. fockei as Lentinus calvescens from Martinique Switzerland is due to spores of P. opuntiae from (see Table 2). Singer (1986) considered P. fockei the Mediterrean region, where it is common. and as a valid species in the subgenus Lentodiellum to­ on the other hand. P. djamor and P. opuntiae are gether with P hirtus. P concavus, and P calyx, rwo :dosely related species, as will be discussed. among others. Recemly, Dickson (personal com­ Other questions will focus on the authenticity of munication) showed the author that Hedger is the identification of the strains. P. lutealbus from studying sorne specimens of P. fockei gathered in Africa (Pegler. 1972, 1977 a) is closely related to Cuyabeno. Ecuador in the postprogram of ::m ex­ P. djamor; it presems yellowish to whitish pileus pedition to Ecuador (Hedger et al., 1995). and pleurocystidia. It is interesting to observe that P. levis seems a valid species, as discussed by P djamor from Cuba is unknown although it is Guzmán (1975), based on Singer (1962) and Far­ surel y a common mushroom there. Kreisel ( 1971) low (1929). Singer (1986) ¡ilso recognized this reported only from Cuba P hinus and P. concavus species. However, Pegler (l98Jbl stated that rhis (see later), and Pegler ( 1988) reported only P mushroom is Lentinus following the po~ition of flavolanatus; however, Pegler (1983a) did not Murrill (1915). P. laciniaro-crenatus is known consider P flavolanatus from. the Antilles. E ven only fromArgentina (Singer, 1950), where ir seems Singer ( 1986) did not consider it. common (Guzmán. 1977b). It is a not a we!l­ P. dryinus is a good temperare species, dis­ known species as irs hyphal system is unknown tinguished by its veil. P. corticatus s. str. is con­ (Singer, 1986). It is an edible mushroom. cultured sidered by Singer (1986) and Kaarik (1992), in thar country, although B lumen fe Id <1994) did among others, as a synonym of P. dryinus, although not consider this species . P. opuntiae is another P dryinus in Mexico seems a complex as will be species with whitish basidioma and whitish spore discussed. P. elongatipes described b~ Peck from print, and sometimes an indeterminate species the ~astern United States is nor a well-known (Hilber. 1982. 1997). It was described from Alge­ species. According to Redhead ( 1986) the cor­ ria on Opuntia (Cacraceae) and it has J. broad dis­ rect name is Hypsizygus tessullatus. Singer tribution in the Mediterrean region on this planr. ( 1986) recorded this taxon as H. tessullatus based and on Agave and Yitcca (Amarilidaceae) and on on Singer and Kuthan ( 1980). Pleurotus albellus Phytolacca (Phytolaccaceae) (Pilát. 1935; Kühner from the Antilles (Pegler. l 983a) seems a closely and Romagnesi. l 953). Dennis (1970) recorded related s p ~tes to P. elongatipes, far the w ell-de­ the species from Venezuela. Pegler l 1977 a) re ­ veloped stipeand white basidioma. Indeed a care­ ported P. opuntiae from Kenya and Naivasha (both ful study is necessary in these mushrooms. in eastern Africa) with the same lecythiform cheilo­ P. hirtus is a neotropical complex. It is known cystidia as those observed in the Venezuelan ma­ from SouthAmerica (Singer, 1956), Cuba (Kreisel, terial by Dennis (1970) and in P. catephes (Pegler, 1971 ), and Mexico (Guzmán, 1975). For Singer l 977a). García-Rollán (1998) in Spain considered (1956) P. hiTtus is composed of four species: this species good for commercial culture. P yuc­ P. hirt;lS S. Str., P. cafvescens,. p concavus, and cae described on a stem of Yucca is a synonym P. calyx. All of them presem white basidiomata. (Pilát, 1935). The reports of P. opuntiae from For Pegler (1983a) P. hirtus s. str. is a synonym of Mexico (Tlaxcala) and New Zealand (Petersen, Lentinus scleropus; P. calvescens is a synonym l 995a,b; Petersen and Krisai-Greilhuber, 1999) of L. striatulus, and P concavus is L. concavus are confused. These authors considered a fungus (also for Comer, 1981 ), and P. caiyx is a synonym identified first as P. agaves. and then as P. opun­ of L. patulus (Pegler. l 983a). Of these names, L. tiae. However, Nicholl ( 1996), as discussed, ob­ striarulus. besides its synonym P caivescens. also served that the New Zealand strains are com­ has the synonym P. fockei (Pegler, l 983a), but Pe­ patible with P djamor. but not with those from gler (l 983b) considered this lacter as a valid Mexico as P agaves (P. agaves "is much smaller"; species, common in the Caribbean region and Petersen. personal communication) or P. opun­ Brazil. and closely related to P. concavus. from tiae. P agaves described on leaves of Agave by

105 Dennis ( 1970) from Venezuela is a related and interbred strains of P. pulmonarius with P co­ poorly known species in this imeresting complex. lumbinus. P. ostreatus, and P. comucopiae, and The Mexican mushro.om of Petersen seems to be described negative results. They distinguished P. cornucopiae s. Guzmán (1977a), also reported P. pulmonarius for its whitish to brownish pale on Agave l.eaves, as Petersen found those mate­ basidioma, P. ostreatus for its brown dark to rials from Mexico. Even Petersen (personal com­ pale pileus. and P. columbinus in its grayish blue munication: 1997) considered fi.rst his Mexican pileus. Romagnesi (1969) separated P ostreatus material as P cornucopiae, but after a mating test from P. pulmonarius and P. comucopiae in the it checked with P. pulmonarius. He observed also color of the spore print and the size of the spores, that this Mexican material was selling in a popu­ besides the hyphal system. P. osrreatus var. ¡?orida lar market at Tlaxcala city, as Guzmán (1977 a, nomen nudum, also known as "P. florida." was 1997) observed with P. cornucopiae in several proposed by Li and Eger ( 1979) from a strain iso­ popular markets of Mexico. As the plants Opuntia lated in Florida (United States). This name was and Agave are of Mexican origin, the distribution never published following nomenclfiture and tax­ of P. opuntiae is obviously in Mexico. Neverthe­ onomic rules. but unfort~nately it qeated great less the species has not been well defined because confusion, because severa! commercial strain~ the reports from Mexico ( e.g .. Estrada-Torres and were used in cultures in tropical countries, such as Aroche, l 987; Petersen and Krisai~Greilhuber. those reponed by Go et al. ( 198 l ), Khanna and 1999) need to be confirrned. Petersen and Krisai­ Garcha (1981 a,b ), Rajarathnam and Bano ( 199 l), Greilhuber ( L999) stated that an epitype is needed. Buswell and Chang (1993), Mirtar et al. ( l993 ), Guzmán ( l 977a) did not consider P. opunriae in and Peberdy et al. ( 1993). Eger's fungus is a syn­ spite of the seven widely distributed species of onym of P. pulmonarius (B resinsky et al., l 97 6; Pleurotus reported from Mexico, but P. cornuco­ Hilber, l982; Singer, l 986: Singer and Harris, piae considered by him is strongly related. P. op­ L987: Watling and Gregory, L989: Guzmán et untiae is close to P djamo1: even Comer ( t98 l ) al.. 1994). Guzmán et al. (1994) obtained assumed close relationships between them. Indeed, basidiomata of P. pulmonarius from a strain ,- a careful morphological. genetic. and biochemical of Eger deposited in Mexico by Zadrazil study on this complex and even on P. levis ob­ through Martínez-Carrera (Martínez-Carrera et served also in Mexico on Agave leaves is neces­ al., 1988a,b). Moreover, the confusion on Eger's sary to define these species. mushroom was more complicated, because Regarding P pulmonarius, it had been con­ Stamets and Chilton (1983) erroneously reponed fused with P ostreatus several times; as Eger et al. this mushroom as P floridanus, which is an ab­ ( 1979) stated that P. pulmonarius is a temperate solutely independent species described by Singer tolerant of P ostreatus. However, Hilber ( 1982), ( 1986). Pleurotus pulmonarius presents white to Bresinsky et al. ( 1987), Cailleux and Joly ( 1993a. whitish or gray pileus, white spore print. and a b); Petersen and Hughes (1993), Vilgalys et al. monornitic hyphal system. Moreover. P. pul­ (1993), Zervakis et al. (1994), and Petersen and monarius is common in summer and P. ostreatus Ridley (1996) separated P. ostreatus from P. pul­ in winter, as observed Cailleux and Joly (1993b) monarius based on their incompatibillty and mor­ and Petersen et al. (l999). Guzmán et al. (l994) phological characteristics. Petersen et al. ( 1999) found that a Cuban strain identified as P. columbi­ separated in their key these two species by the dif­ nus (IE-136), obtained from a Czechoslova­ ferent colors of the pileus and by the season when kian institution, thp.t produces blue basidiomata, they develop their basiomata, winter for P. ostrea­ is compatible with the Zadrazil strain. This tus and spring and summer for the latter, in spite . shows how problematic it is to base the taxonomic of the fact that Petersen claimed severa! times decisions only on the interbreeding. P. pulmonar­ that the mating tests are the most important meaos ius is common in the Mediterranean region and in of idemification in Pleurorus. as discussed later. the eastern United States as stated previously B unyard et al. (1996) separated both species on and observed by Singer and Harris ( 1987) and the basis of DNA studies. Bresinsky et al. (1976) Guzmán (1996).

106 P. sapidus seems conspecific with P. ostreatus cluster indicative of a genetic affinity. Far Chang (Hilber, 1982: Zervakis et al., 1994) or wirh P. pul­ (1991) the commercial cultures of P. sajor-caju monarius (Zervakis and Balis, 1995). However, belong to the complex P. ostreatus. Petersen et al. for sorne authors (e.g., Peberdy et al., 1993) it is a (1999) observed that P. sajor-caju is a misidenti­ valid species. Buchanan (1993) considered P. fication of P pulmonarius. sapidus conspecific with P. comucopiae. Pleuro­ P. tuber-regillm considered by Singer (1949. tus sajor-cajll is a southern Asiatic and African 1962. 1975, l986) belongs to Lenlinus according species, with a well-developed annulus at the to Pegler ( 1972, 1977a. l983b, 1986) and Watling apex of the stipe, and considered by Pegler (1969, (1991, 1998), or to a Panus according to Comer 1977 a, l 983b. 1986) in lentinus. However, the ( l 981 ). This mushroom, in fact. presents a cinna­ cultivated commercial strains of P. sajor-caju in mon brown pileus and stipe (Pegler, 1972; Watling, Europe, Asia. and the United States belong to 1998) not common in Pleuroms. and its anamorph P. ostreatus or P. pulmonarius, based on the ba­ state is known as Pachyma (Singer, 1986). It is known fromAfrica and Asía . .and distinguished bv sidiomata obtained (Guzmán et al., 1994: Pegler -... '- ... et al. , 1999). P. .wjor-caju is known among the its large buried sclerotiU:m up"io 20-30 cm long. commercial edible mushroom products as "gray Recently, in a molecular study Neda ano Nakai oyster mushroom" or "phoenix-tail mushroom."' (1995) found thar P. tube r-regium is distantly re­ Those strains of Chang and Hayes ( 1978), Nair lated to Lenrinus and Panlls, and they concluded and Kaul ( 1980), Hilber ( 1982. 1989), Mueller and that this mushroom belongs to Pleurows. However. Gawley (1983), Quimio (1986), Martínez-Carrera Petersen et ::i.l. ( 1999) pointed out that micro­ et al. (l 988a,b ), Chang and Miles (1989), Okwu­ droplets were fo und in cultures of this fungus jiako (1990). Aslan-Azizi et al. (1990), Zervakis (Hibbett and Thom, 1994 ), which are typical for and Labarere (1992), Yoo and Cha ( 1993), Chang Pleurotus. ( 1993 ), Chang et al. (l 993a.b) ::md Gibriel et al. P. se rotinus is Panellus se rotinus according \ ( 1996) surely belong to P. ostreatus or P. pul- to S inger ( 1949, 1962. 197 5, 1986) or Sarcomyxa monarius for the flaveliform, not infundibuliform serotina according to Horak (1 968), although it basidiommata developed and without annulus. was considered in the bibliography as Pleurorus The strains of '' P. sajor-caju'' in Mexico (IE-44 or (e.g .. Rea. 1922) or ;is Crepidorus (e.g., by Mur­ INIREB-49) used by Martínez-Carrera et al. rilL 1916). Another indeterrninate confused species (l 988a,b) developed P. ostreatus basidiomata reported in the bibliography is P. komamitzkyi. con­ (Guzmán et al. , 1994). The basidiomaca obtained sidered by Hilber ( 1982) as having uncenain tax­ by Mueller_:ind Gawley (1983) agree wirh P. os­ onomic position. but far Singer (1986) ir is prob­ treatus or P. pulmonarius. Kurtzman and Zadrazil ably a Clitocybe. Hilber (1997) considered this a ( 1982) stated that the strains ITCCF l 725 and valid species from Asia. P. spodoleucus is another ATTC 32978 identified as P sajor-caju and cul­ not well known species, sometimes considered as tured in Asia produce excellent mushrooms both a synonym of P. osrreatus (Imazek.i and Hongo, in flavor and in texture, unlike P. sajor-caju. 1983). Singer ( 1943) consÍdered ita valid species. Smith .( l 993) reported strains of P. sajor-caju in but Singer ( 1986) did not consider this species. A the Intematíonal Mycological · Institute at Kew genetic study on this mushroom was made by Yoo without any information. A good color figure of and Cha (1993). May and Royse (1988) and Ma­ the basidiomatae of P. sajor-caju is found in Pe­ gae and co-workers in 1990 (in Buchanan, 1993, gler (1972). Hilber (1989) assumed that those re­ who conside.red this name as P. ostreaws) studied ports of P. sajor-caju with synnema stage (Nair the enzymes of this mushroom. Benjamín ( 1995) and Kaul. 1980) belonged to P cystidiosus. From reported sorne medicinal properties of this species. another point of view, Zadrazil and Kurtzman P dubius is a not well known species. It was con­ (1982) and Roxon and Jong (1977) considered sidered by Re id ( l 962) from New Zealand based on P. sajor-caju as a valid species of Pleurotus, but · Stereum dubium described by Lloyd in 1925. But Zervakis et al. ( 1994) found that the strains of this mushroom probably belongs to another genus P. sajor-caju form with P. pulmonarius a tight (Hohenbuehelia'?), bec::mse it presents cystidia with

107 thick walls and subglobose spores. P subareolams tures and also the hyphal system in any of the is another poorly kn.own species, described from three species. Concerning use of only interbreed­ the eastern United States. According to Anderson ing as a method, Petersen and Ridley (1996) et al. (1973) and Groves (1962) it is el ose to P. os­ showed that a strain from New Zealand named P creatus. but the spore print is white. Probably it is pulmonarius, is compatible with P. ostreatus. P. P. pulmonarius. eryngii, P. populnus, P. pulmonarius, and P. abi­ eticola, but not with P. djamor from Mexico. More recently, Petersen and Hughes ( 1997) NEW METHODS AND PROBLEMS stated that P. abieticola, described from Russia. IN THE lDENTIFlCATION OF THE has negative interbreeding with all the species SPECIES OF PLEUROTUS cited above. The observations of Reid et al. (1998) on P. fuscosquamulosus, that this species Modern methods based on genetic and bio­ is compatible or not (0% or 100%) with P. cys­ chemical studies seem to be overemphasized. tidiosus according to the origín,_of the strain. Moreover. the authors who follow them claimed showed also. together with the Vilgalys ;ind Pe­ that the morphological taxonomic features have tersen observations discussed above, how co,n­ been found insufficient in Pleurotus. Buchanan fusing the mxonomy of Pleurotus is when based (1993) stated that macroscopic feamres in isola­ onl y in interbreeding tests! tion are not very useful to support taxonomic con­ The modern concept of the species of the clusions and the commerciai stro.ins may c:1rry genus Pleurows needs to deal with morphological ambiguous or incorrect names. Petersen (1995b) features. such as form of basidioma, type of sur­ claimed that sorne taxonomic conclusions based face of the pileus and stipe. :md color of all the on enzyme studies on P!eurows ;ire inexacr. be­ parts, including the spore print and the type of cause thev did not include inrerbreeding exoeri- hyphal system. besides biochemical and molecular • - 1 ' ments. such o.s those of .\fay and Royse (1988) data. and interbreeding. It is necessary to revise and Zervakis et al. ( l 994). in which even P sajor­ the status of severa! species. and then the number caju is considered a valid species. even though of species of P!eurorus. Petersen (l 995b) and Pe­ this is an erroneous name for sorne strains of P tersen et al. ( 1999) claimed that it is necessary to ostreatus or P pulmonarius. Many modern au­ make a contraction, but if we base the contraction thors (e.g .. Vilgalys et al.. 1993: Egeret al.. 1979) of thé species only on mating tests (as severa! au­ claimed that the macroscopic and microscopic thors claimed in other mushrooms, e.g .. Boidin, characteristics a?é-unreliable for classification in 1986) or on biochemical or molecular studies. we Pleurotus (e.g .. in the P ostreatus complex). can make faxonomic mistakes. Certainly, the inter­ However, Vilgalys et :il. ( 1993) distinguished breeding and biochemical studies play an impor­ three groups of P ostreaws in the U nited States: tant role. but not the only one. Indeed the problem (l) P ostreatus s. str. with lilac-gray to purplish is complex. because first we need to have an ex­ vinaceous spore ppnt in the east, even in Florida act concept of the species, as Petersen and Krisai· [where Li and Eger ( 1979) describec;l their "Pleu­ Greilhuber ( 1996) presented for P. ostreatus. rorus florida." actually contaxic with P. pulmon­ The almost "common" taxonomic confusions arius or P ostreatus. according to the strains], on Pleurotus have opened the door to genetic (in­ and also from California and Arizona: (2) P. pu.l­ terbreeding), biochemicaL and molecular studies monarius from the northern states and also from (e.g., Eugenio andAnderson, 1968; Blaich. 1976; California and Arizona. with a white. yellowish, Eger, 1978; R:aper, 1978; Volland-Nail et al., buff to lavender gray spore print ::md with a dis­ 1981; Hilber. 1982: Bresinsky et aL 1987; May tinctive coloration in the basidioma. contrary to and Royse, 1988; Royse and May, 1993; Vilgalys the European interpretations. which reponed this et al., 1993, 1996; Ogawa, 1993: Peberdy et al., species as whitish. lacking darker pigments; and 1993; Buchanan, 1993; Vilgalys and Sun, (3) a new species, named P populinus from the l 994a, b; Labarere and Ira~abal, 199 5; Zervakis northern states with whitish. not lilac spore print. and Balis. 1995; Bunyard et al.. 1996: Müller, They did not submit information on other fea- l 997). The variation in the form ;ind color of the

108 basidioma of Pleurotus in culture is so large that tests are necessary to elucidate the specíes. Eger it often leads to wrong identifications or to inap­ (1978) and Eger et al. (1979) established that the propiare conclu.sions. It is known that P ostreatus, definitions of species. especially in the genus Pleu­ P. djamor, P. pulmonariu.s, and others present in rotus, are controversial. They based their concept culture sorne features not typical in the wild ba­ only on interbreeding experiments and found that sidioipa, for example, the presence of long stipe. several different species are conspecífic, but they deformation in the basidioma similar to synnema, used strains not so well identified, as stated by May change in the color. etc., such as those observed and Royse (1988). Eger "described" P florida or by Zadrazil (1974, 1978), Desbiens et al. (1978), P. osrreatus var. j lorida. a very questionable name, Zadrazil and Kurrzman ( 1982), and Guzmán et al. as it was díscussed. Vilgalys eral. (1996) wrote: (1993a, 1994). The cultivated strains and inter­ "Mating compatibility studies and molecular breeding among them. and the biochemical and phylogenetic analysis provide a useful framework molecular studies in Pleurotus, without a careful · for understanding specíes concepts and taxonomy taxonomic analysis, are creating an atmosphere of of ... Pleurorus." They stÚdi~d 15 groups associ­ confusion on the taxonomy of the genus. Neda ated wüh one or more morpnological species, and Nakai (1995) and Perersen et al. (1999), for among them P. agaves from Mexico (a cenfused example, determined thar Pleuroms ruber-regium, complex related to P. cornucopiae sensu Guz­ discussed previously as Lenriríus or Panus, pres­ mán. as previously discussed) and others from New ents more affinities with Pleurotus than with those Zealand. Australasia. Brazil. and P levis. this lat­ genera. Also Neda and Nakai (1995) showed that ter previously considered as P dryinus in Vilgaiys P. ostreatus and P. pulmonarius, two different and Sun (1994a.b). After interbreeding and bio­ species, are very close and they did the same with chemisrry studies in a strain from Quebec (Crn­ P. abalonus and P. cvstidiosus. Zervakis and Balis ada), Volland-N ail et al. (1981) could not define (1995) discussed that P. sajor-caju i~ an Asiatic the species. variety of P. pulmonarius. Bresinsky et al. (1987) Petersen ( l 995b) rejected taxonomic analyses .( derermined that mating and fruiting are two im­ based only on morphological features. He stated: portam sreps to be followed in the identificarion ·'Morphoiogícal characrers of basidiomata are !ike of Pleurocus and rhey claimed P. ostreatus plays the tip of an iceberg:· However, later. in Petersen an important role in the specialization of several et al. ( 1999), he distinguished in a key 15 species species. However. wirh P. cystidiosus and others. of Pleurorus, based only on morphological and Bresinsky et. al. (1987) said rhat the "compatibil­ color features, even on the seasonal fruiting. It is ity is reducect::.__( P ostreatus and P cystidiosus are necessary to be cautious also in those cases of two different species ! ). These authors stated that hybridization by protoplast fusion. For example. between P. osrrearus and P pu.lmonarius genetic Ogawa (1993) hybridized P. cornucopiae with barriers were erected early in the separation of Lentinula edades and Lyophyllum decastes (Fr.) these fungi. Sing .. Regarding the species concept, it is generally It is certainly necessary to revise several de­ accepted thar species are not .capable of inter­ scribed species and also to describe new species, breeding, or the hybrids between them are sterile on the basis of modern methodology. However, or rarely formed in nature, as stated by Boidin in this case a careful concept of a species has to ( 1986) and Raven in 1994 (in Nature Conservation be followed. Hawksworth (2000) commented on 44, taken from Petersen. 1995b) in several fungi. the concept bf species according to Petersen and Bresinsky et al. (1976) claimed that the species Hughes (Í999), who stated that the species con­ concept should be based primarily on macro­ cept wíll be based on evolutionary and reproduc­ and microscopical features. on macro- and micro­ tive information, but they concluded that a species chemical characters, and on ecologícal observa­ is what a good taxonomist recognizes; in other tions. However, they observed that physiological words, a species depends on the investigator cri­ and biochemical information would also be use­ reria. Hawskworth observed well that these Pe­ ful to understand the variation of the species. For tersen and Hughes ideas reflect the apocryphal Hilber (1982) and Boidin (1986) interbreeding concept of the species. This is absolutely true.

109 DESCRIPTION OFTHE GENUS TABLE 4 PLEUROTUS ANO ITS A Methodology to ldentify the Species TAXONOMIC POSITION of Pleurotus (The numbers are in arder of importance.} To define the limits of Pleurotus, the concept 1. A careful study of the basidioma (color of ali the of the gerfus accepted in the present article is parts, texture, type of surface of the pileus, position shown in Table 3. This concept is an adaptation of and surface of stipe, presence of veil) that of Fries (182 l. 1838), Pegler ( 1977a, 1983 a. 2. Color of the spore print in fresh 1986). Singer (1 986). Batisra-Pereira (1988), 3. Wild substrate Hilber (1 989. 1997), Bas (1990), and Kiiarik 4. lsolation of strains and their interbreeding with other strains (1992), among others. Both monomitic and 5. Obtaining fructifications (basidiomata) of the dimitic hyphal systems are considered in the cultured mycelium genus. and the comext is íleshy or soft. No metu­ 6. Biochemical studies loids are present in the genus but pleurocystidia 7. Molecular studies '· are common in sorne species (e.g., P cystidiosus). 8. A careful correlation of the above poi"ts The surface of the pileus is generally smooth, al­ though scaly in so rne species (e.g .. in P cysridio­ sus and P smithii), rarely viscid (e.g .. in P vis­ repon ed fragranr. sweet. and floral aromas fo r P cidus), or most frequently dry. The color of the euosmus; slightly fragrant. mushroom. fruity, bir­ spore print presems a wide variation from whire. ter cocoa. anise. almond. and yeast for P ostrea­ yellowish, gray, lilaceous to pink. The ílavor is tus: and fragrant fo r P sapidus. usually fu ngoid and pleasam. sometimes like fi sh However. fo r a correct identification of the or anise. P djamor presents both odor and t1avor species of Pleurotus, it is recommended to obtain that are farinaceous. Jong ;,ind Birmingham O9 93) a strain of the material and to obtain basidiomara and try interbreeding the strain with others fro m a we il identified species to obtain fertile basidio­ mata and to study the developmem of these latter. TABLE 3 B iochemical and molecular smdies will be useful Description of the Genus Preurotus in this taxonomic srudy. But as shown in Table 4 and Fig. 1, it is very imponant to fo llow morpho­ Basidioma flabelliform or sometimes infundibuliform. logical and color features, genetic studies. and Pileus white, whiti~t:li_ yellow, brownish pale or dark, grayish brown, blue, grayish blue, pink or orange pink, biochemical and molecular research. Further­ dry or sometimes viscid or .hygrophanous, smooth or more, point 8 in Table 4 and Fig. l will be the scaly, or tomentose at the center. Lamellae decurrent, most important in the taxonomic smdy of Pleuro­ uniformly white, whitish, pink or orange-pink, with con­ tus. but the identification should never be based ex­ colorous edge. Stipe absent or as lateral or eccentric, clusively on genetic, biochemical. or molecular short or long, smooth, tomentose, or hirsute mainly at the base, solid. Velum and ring present in sorne studies, without the information of poínts l and 2. species. Confext fleshy or soft, white or.whitish. Odor Conceming the taxonomic position of Pleu­ and flavor pleasant, fungoid , or farinaceous. Spore rotus. it is considered here as a member of the print white, yellowish, pink, grayish, lilaceous, lilac­ Family Pleuroraceae, together with Lentinus Fr., gray, or pale vinaceous. Spores hyaline, cylindrical, thin N eolentinus Redhead et Ginns. and Lentinula Earle walled. Pleurocystidia absent or present, never as in Order Poriales•(or Polyporales). This position metuloid. Cheilocystidia usually present, but not as metuloid. Hyphal system monomitic or dimitic, with in­ follows in part Singer (1 986) and takes into con­ flated hyphae up to 25 µm wide. Hymenophoral trama sideration those observations by the author (in Guz­ subregular or irregular. Pileal surface undifferentiated. mán et al.. 1997) on Lentinula. This latter genus Clamp connections present. Mycelia white, with nem­ is considered by Pegler ( 1983c) as a member of atode-trapping structures. Anamorph state as syn­ the Tricholomataceae family in Tribe Collybieae nemae with black globose heads and black spores. Habitat on living or dead deciduous plants (mainly in Agaricales, on the basis of the inflated hyphae trees), rarely on coniferous trees, growing also on sev­ of the context. However, as observed by Comer era! lignocelulolitic substrates. (1 98 1) and Guzmán et al. ( 1997), those inflated

110 hyphae of Lenrinula are like [hose observed in 1975). Recently, Sobal et al. (1997) developed in Pleurotus, but at the same time completely dif­ the laboratory basidiomata of this species with a ferent of those .of the Tribe Collybieae in the Tri­ veil only in the early stages. The report of Gándara . " cholomataceous fungí, for example, Collybia (Fr.) (1930) for P eryngii, without any information, Staude. Collybia presents the inflated hyphae with seeÍns to be a rnistake, because this species is un­ thin walls, 0.5-1 µm thick. while in Pleurotus and known in America. P bajocalifomicus is known Lent(nula the walls are 1-2 µm [bick. In conclu­ only in a deserr from Baja California Península sion, there are not subsrnntial differences between (Moreno et al. , 1993). The spore print was not Pleurotus and Lentinus vs. Lentinula to consider recorded; it presents a monomitic system, colly­ these genera in different orders. bioid habit. and no veil. For P catephes see ear­ lier. Concerning P. elongatipes, it was reported without any description from a tropical area of THE KNOWN SPECIES OF PLEUROTUS Mexico (Welden and Guzmán, 1978). A careful IN MEXICO AND THE!R CULTIVATION study of those materials of tn.is mushroom is nec­ essary at ENCB. This species"seems related to P. There are 23 species and varieties of Pleuro­ albellus (Pegler, l 983a) from the Antilles. P cor­ ws reported from Mexico (Table 5), of which nucopiae was reported by Guzmán ( 1977a) grow­ only 7 seem to be valid raxa. Of these, Pegler ing on the base ofleaves of Agave (mainly A. arro­ (1983b) considered P hirtus and L. levis as be­ virens Karw.) in the central plateau of Mexico. longing to Lentimts (L. scleropus and P levis. re­ This species seems to be the P. opuntiae sensu Pe­ spectively). P. hirtus is common in the tropics. tersen (Peterse n and Krisai-Greilhuber, 1999) or while P. levis is found in the subtropical humid P. levis according to Guzmán (1977a) as dis­ forests and also in temperate fo rests (Guzmán. cussed. P /loridanus was reported by Herrera

TABLE 5 The Known Species of Pleurotus in Mexico (Only the first bibliographic reference of the taxon is presented in each case. Names in boldface are recognized species.)

P agaves (Petersen and Ridley, 1996) (? P opuntiae) P. bajocalifornicus (Moreno et al. , 1993) (only known from Baja California Peninsula) P catephes (Guzmán, 1983) P cornucopiata (Q_uzmán, 1977a) (confused with P opuntiae and P levis) P. djamorvar. djamor (Guzmán et al.. 1993a) P djamor sensu Petersen and Hughes (1993) and Petersen and Ridley (1996): ? P opuntiae P djamorvar. roseus (Guzmán et al., 1993b) (P djamorvar. djamoi) P djamorvar. salmoneostramineus (Guzmán et al. , 1993b) (P djamorvar. djamof) P dryinus (Guzmán, 1975; Valenzu.ela et al., 1981) (a complex species in Mexico) P elongatipes (Welden and Guzmán, 1978) P eryngJi (Gándara, 1930) (an erroneous report?) P eugrammus (Welden and Guzmán. 1978) (Nothopanus eugrammus) P flabellatus (Welden and Guzmán, 1978) (conspecific with P djamof) P f/oridanus (non P florida s. al.Jet.); ?Hohenbuehelia (Herrera, 1960 from Isla Socorro) P. hirtus (Guzmán, 1975); Lentinus sc/eropus P hygrophanus (Guzmán, 1983) (Nothopanus hygrophnus) P. levis (Guzmári, 1975); Lentinus /evis P mexicanus (Guzmán and Johnston, 1974) (conspecific with P djamor var. djamoi) P. opuntiae (Estrada-Torres and Aroche, 1987); still poorly known species related with P agaves, P cornucopiae, P djamor and P levis P ostreatoroseus (Welden and Guzmán, 1978) ( P djamor vp.r. djamof) P. ostreatus (Guzmán, 1977a) (unknown in the wild) P roseopileatus (Guzmán, 1977a) (P djamorvar. djamof) P. smithii (Guzmán, 1975); anamorph: Antromycopsis guzmanii (A smithi1) (Guzmán et al., 1980; Stalpers et al., 1991)

111 (1960) from Isla Socorro (in the Pacific Ocean), TABLE 6 with two good picrures, bur without any descrip­ Commercial Cultivated Species of Pleurotus tion. It is necessary to check the herbarium mate­ in Mexico rial at MEXU of Herrera·s specimens; but as dis­ P citrinopileatus cussed this · Singer's mushroom may be long to P columbinus Hohenbueheiia. This report is independent of those P dja,mor var. djamor cultures referred to as P. f lorida s. auct. or erro­ P djamor var. sa/moneostramineus ( P djamor var. neously nafu.ed P. jloridanus (see earlier). P. djamoi'j smirhii (Guzmán. 1975; Guzmán et al., 1980, P djamorvar. roseus (P djamorvar. djamoi'j 1991; Mora et al., 1984) is fairly common in the P ostreatus P pulmonarius ( P florida) temperate regions (e.g., Valley ofMexico City) and P sajor-caju s. auct. (P ostreatus) (not Lentinus subtropical regions (e.g., Xalapa and Morelos). It sajor-caju) has been observed as parasitic on trees: Schinus molle L., Populus spp .. Quercus spp .. and Psidium ' guajava L., although in Cuba (Rodríguez and thors also recorded P comucopiae. P"opuntiae is Camino, 1990) i t was reported from a Ficus re tusa a species not well known in Mexico. The recor~ L. As discussed. P smirhii is close to P cystidio­ of Estrada-Torres and Aroche ( 1987) in Mexico sus, but differem because of the pleurocystidia needs revising, even that by Petersen and Krisai­ present in the laner. although no interbreeding be­ Greilhuber ( l 999) gathered in Tlaxcala on Agave tween bmh species has been done. It is interesting leaves. P. opunriae sensu Petersen seems close to m observe that P. smithii in culture frequently P. cornucopiae or P levis (both sensu Guzmán. produces the synnematous phase (Guzmán et al.. l977a). P. ostreatus s. str., following Petersen and 1980; Spinedi, 1995). Mora et al. ( 1984) described Krisai-Greilhuber (1996), is either unknown in the wild synnematous phase growing on the stipe Mexico or perhaps grows in coniferous and tem­ 1 of a wi.ld basidioma, ::md \'Iartínez-Carrera et al. perare forests in the north of the country. P dryi­ ( 199 lb) obtained basidiomata from a differemia­ nus is known only in the northem and central ; tion of small synnemata. states of Mexico (Guzmán. 1975; Valenzuela et There is confusion regarding the presence of al., 1981 ). It was recently reported by Nava and P. ostrearus in Mexico. It seems that all the reports Valenzuela ( 1997) from the State of Mexico. The of this species in Mexico (e.g. , Guzmán. 1977a, well-developed veil, as well as the size qf spores, 1983: Martínez-Carrera. 1984: Martínez-Carrera are the important features of this species. How­ and Morales. 198'8; Acosta-U rdapilleta et al. , ever. P dryinus seems a complex, according to the 1988. 1995: Bemabe-González and Garzón-Mayo. size of spores, at least in Mexico, that is now in 1995: Herrera and Ulloa. 1998) be long to P study by the author in collaboration with Valen­ Jjamor, as observed by Guzmán et al. (l 993b ). zuela. Navarro et al. ( 1996) found that Mexican strains Concerning the commercial and/or experi­ of P. ostrearus frorn Morelos were incompatible mental cultures on Pleurotus in Mexico, 8 ra·xa with strains from the United States aqd Gerrnany. (Table 6) are the most commonly considered. Of P. djamor is the "tropical form" or the "white these, P. ostreatus, P. columbinus, "P. sajor-caju," form" of P ostreatus, as discussed by Kaul and "P florida, " P pulmonarius, and P djamor (this Kachroo ( 1970) in India, and considered by latter with its rypical and pin.k forms) are the most Singer (1956). P. djamor is the most important important, all of them are foreign strains (from the species in Mexico for its wide tropical and sub­ United States or,European) (except far those of P tropical distribution (Guzmán et al. , 1993b). P. djamor). Nonetheless, only P ostreatus is com­ mexicanus (Guzmán and Johnston, 1974) is con­ mercially cultivated using high technology by specific with it: however. Guzmán et al. ( 1993 b) re­ two or three imponant factories (Martínez-Car• poned a confused monomitic hyphal system in the rera· et al. , 199 la). But many small companies not so well preserve type at ENCB. It is probably cultivare Pleurotus in small mushroom farms. that this hyphal system is really dimitic. Herrera These cultures began in the region of Xalapa in and Ulloa ( l 998) recorded P. mexicanus. These au- the l 980s and now are spreading in many cities of

112 Mexico (Mata, 2000). They use mainly straw, but with Bernabe-González as head; and (7) Univer­ sometimes coffee pulp, sugar cane bagaze. tequila sity of Morelos in Cuernavaca. with Acosta-Ur­ bagaze (from Agave tequilana L.), and other agri­ dapilleta as head (e.g .. Guzmán-Dávalos et al., culture wastes (Martínez-Carrera, 1984; Martínez• 1987, 1989; Acosta-Urdapilleta et al., 1988, 1995; Carrera et al.. 1984, 1988b; Guzmán-Dávalos et Soto-Velazco et al. , 1991 a,b; Martínez-Carrera et al., 1987; Acosta-Urdapillera et al., 1988; Guzmán­ al.. 199 la,b; Ramírez-Carillo et al.. 1991: Guzmán Dávalos and Soto-Velazco. 1989; Soro-Ve!azco et al. , 1993a; Mata and Gaitán-Hemández. 1995; et al., 199 la,b: Cedano et al., 1993; Guzmán et al., Bernabe-González and Garzón-Mayo, 1995; 1993a; Bemabe-González. and Garzón-Mayo, Hernández -Ibarra et al. , 199 5; Leal-Lara in Pare­ 1995; Mara and Gaitán-Hernández, 1995; Nieto­ des et al. , 1996; Sánchez in López-Arevalo et al., López and Sánchez, 1997). Aguilar et al. (1993) 1996; Nieto-López and Sánchez, 1997). Sorne of presented an interesting rural experiment. where the few chemical studies on Ple uro rus developed a commercial farm for cultivating P ostrearus was in Mexico are those of Trig?s and Martínez-Car­ set up in an Indian region of Mexico (Cuerzalan rera ( 1992) and Trigos et al. ( 1994) and are based and Puebla). Sobal et al. (1997) discussed the on the coment of ergosterol. " Mexican production of Pleurotus in Mexico, 356 tons in 1990 and 1825 tons in 1997. versus the world production of 909 .000 to ns in 1990 TRADITIONAL MEDICINAL USES (Chang and Miles, 1991). The center of Xalapa OF THE SPECIES OF PLEUROTUS (first as INIREB, and actually as Instituto de ANO TREATMENT OF ALLERGIES Ecología) was the firsr insritution in Mexico to develop experimental cultures in a special mush­ Al though they are very common in Mexico room house in l 983 (Martínez-Carrera. 1984: compared to other edible mushrooms (Guzmán. Martínez-Carrera et aL 1984. i 988a,b; Guzmán 1977a), all the species of Pleurotus have tradi­ et al.. 1993a; Lara-Herrera et al.. 1998). The tional edible and medicinal uses (G uzmán, l 994a.b, commercial srrains of P sajor-caju used (requemly. 1997). However. these mushrooms are so importam as discussed, are P ostreaws or P pufmonarius in the coumry that there are almost 100 common according to the strains. P djamor with its vari­ names for rhese species (Guzmán, 1997), as shown eties (ecological forros. Guzmán et al.. 1995) are in Table 7. Man y of these names are related to the from native Mexican strains. except the srrains habirat. such as those of "hongo'' (= mushroom) of P djamor var. salmoneostramineus. which ir or "oreja"(= ear) "de cazahuate." " .. . de encino," seems is fromA~ia, also rhat of P citrinopilearus. " . .. de izote," " ... de maguey;· " ... del café," The commercial cultivation of Pleurotus in Mex­ " . .. del bagazo de la caña," which mean fun gus ico started in the l 970s (Martínez-Carrera et ::il. , of the tree lpomea spp., of Quercus spp. , of Yucca 1991 a) with European strains of P ostreatus. The spp., of Agave spp., of the coffee pulp, and of commercial product was called "se~a " (plural sugar cane bagaze, respectively. Those common "setas," a Spani.s h name that means mushroom), names together with the word "nanácatl," mean­ and now .rhis c.ommon name is t.Qe mosr used in ing mushroom, are indigenous from the Nahuatl the country for species of Pleurotus. language. The commercial name for the cultivated At present, the following Mexican instirutions Pleurotus is seta, as previously discussed. have research programs on the culture of Pleuro­ As for medicine, the species of Pleurotus are tus: (1) Colegio de Posgraduados at Puebla, with recommended •in Mexico to reduce cholesterol Martínez-Carrera as head; (2) ECOSUR in Tapa­ levels; to figfit diabetes. high blood presure, nerv­ chula, Chiapas, wirh Sánchez as head; (3) Insti­ ous disorders; to promote good memory; as an tuto de Botánica in the U niversity of Guadalajara, antiparasitic; for sexual dysfunction; for rejuve­ with Soto-Velazco and Guzmán-Dávalos as the nation; as a laxative: to darken hair: and as an principals; (4) [nstituto de Ecología in Xalapa, intestinal antiinflammatory and for intestinal ulcer. with Mata as head; (5) Instituto de Química in Small fragments of dry specimens and powder ob­ the University of Mexico with Leal-Lara as head; tained from dry specimens are used. Capsules are (6) U niversity of Guerrero, in Chilpancingo. even made with the powder (Guzmán. 1994a,b).

113 . - TABLE 7. ~ommon Names of Pleurotus spp. in Mexico

Ahuananácatl Mazajielle de agua Cazahuananácatl Menanácatl Cazahuate Mezonanácatl Cazahuatem Me zorras Cuahuiztacnanácatl Oreja Cuauhiztac Oreja blanca Cuauhzahuáíic Oreja colorada Cuazahuananácatl Oreja de burro Cui moni Oreja de cazahuate Chhó xuni Oreja de cazahuate rosa Chiltizcatl Oreja de izote Chilla nalhat Oreja de jonote Hongo blanco Oreja de maguey Hongo blanco de mayo Oreja de palo Hongo de cazahuate Oreja de patancan Hongo de chonotl Oreja rosada Hongo de cocohuite Orejas Hongo de encino · Pechuga Hongo de izote · Pechuga de maguey Hongo de jonote Pleuroto Hongo de la paja Pleurotus Hongo de la pulpa del café Pobnec Hongo de madroño Repollo Hongo de maguey Riruchi Hongo de palo Sacita Hongo de palo blanco Sacocox Hongo de palo mulato Sakitah Hongo de tejomite Seta Hongo de xonote Seta de cultivo Hongo del bagazo Sinche Hongo del bagazo de la caña de azúcar Tasnara Hongo del caié Techalonanácatl Hongo del madrecacao Tepetomananácatl - Hongo del maguey Tepetomananágatl ,.___ Hongo del naranjilla Tetecuin lawakamazlat Trompa de palo lczonami.gatl Trompetitas de los palos lztacnanácatl Tua tasnara Jetch Utuxa yekua Jonacate Xinche Kewaru kowaru Xonanácatl Kikínche Xonocote Kju wada Xonocuahnanácatl Lawakamazlat Xononanácatl Magueyero Xonotnanácatl Malhat kiwi Xumpililomazlat Matomananácatl Yehyecañanágame Yehyegannágame Note: Nanácatl means mushroom in lndian; hongo (in Spanish} means mushroom; cazahuate is the common name of a tree; maguey is Agave; maguellero means that which grows on the leaves of Agave; oreja means ear; pechuga means breast; repollo means cabbage; seta means mushroom; xonote is a wild tree.

114 TABLE 8 Medicinal Properties of and Medical Conditions Treatable with Pleurotus spp. According to Diverse Bibliographic Sources (See text.)

Anticarcinogenic Diuretic Activator of cellular immune system Fever Antibacterial Gastrointestinal disorders Antibiotic Good memory Antiinflammatory Headache Anti tumoral Hematological effects Antibacterial High blood pressure Antiparasitic Hyperlipidemia Antifungal Hypocholesterolemic activity Aphrodisiac (sexual power) Hypotensive renal effeqts Arteriosclerosis (prevention) Intestinal ulcer ' Assist in recovering from fatigue Intestinal antiinflammato~ Asthma Nervous disorders Cardiovascular effects Rejuvenation Constipation Renal effects Cholesterol (reduce) Sexualdecadence Cold Stomach ache Darken hair Vigor (produce) Diabetes

In other countries, severa! spocies of Pleuro­ later. Watling ( 1998) found basidiomata in the - tus have been reponed in traditional medicine or same sclerotia. Gunde-Cimerman ( 1999) and ' they are being exp!ored extensivel y for their phar­ Wasser and Weis ( 1999) discussed the imporrance maceutical utility (e.g. , Buswell and Chang, 1993: of Pleurotus spp. in medicine, their nutritional Sani and Atri, l 999; Gunde-Cimerman. 1999: value. and their importance as a source of severa! Wasser and Weis. 1999). In traditional medicine substances of medica! interesr. they are being used to prevent or assist in more In reference ro the anticancerous activity re­ than 30 d!seases or disorders ¡_ Table 8). Regarding ported for Pleuroms by the above authors. Ben­ the medi"Cinal properries reported in the bib!iog­ jamín ( 1995) discussed, based on the bibliogra­ raphy, care is necessary when referring to the old phy, that P. spodoleucus possesses this activity. books. because the identificarions of the mush­ In P sajor-caju, Tam and co-workers in 1986 rooms can be erroneous. For examp!e, the "phan­ (Saini and Atri, 1999) commented that an aqueous somb ::i" mushroom from India, which belongs to extraer has been reported to reduce the rates a Phellinus. was erroneously identified as P os­ of nephron deterioration in persons suffering treii.tus (i.mpublished notes· of Vaidya in India). from renal failure. Buswell and Chang (1993) Stamets (1993) mentioned P citrinopileatus that recorded, based on Vogel and co-workers, "potentially cures pulmonary emphysema" accord­ Ikekawa and co-workers, Bobek and co-wokers, ing to Chinese sources. Oso ( 1977), Singer and Tam and co-wokers. antitumoral effects, (1986), Watling (1991 , 1998), and Benjamín hypoctioi~sterol effects, and hypotensive activity (1995) discussed the sclerotia of P tuber-regium of extracts of P ostreatus. Chang and Miles that are used by the native peoples in Africa as ( 1989) pointed out the hematological effects food :md for diverse medicinal purposes, such as and cardiovascular or hypotensive and renal ef­ healing stomach ache, constiparion, fever. and. fects of P. ostreaws and P sajor-ca}Lt. Amibiotic high blood pressure. It is interesting that Watling effects and antitumoral effects were discussed by ( 1991 ) reported a sclerotia gathered in Cameroon Cochran ( 1978). ( Africa) and sowed in the tropical greenhouse of The species of PleLtrotus in farming incite an the Botanic Garden at Edinburgh. Seven years allergic reaction or picker's lung in sorne mush-

115 room house workers (Eger, 1978; Zadrazil, 1978; G. Mata, D. Salmones, R. Gaitán-Hernández, Zadrazil and Kunzman, 1982; Chang and Miles, S. Chacón, L. Montoya, and V. M. Bandala, and 1989; Mittar et al., 1993), as observed by the his assistants F. Ramírez-Guillén and F. Tapia, author in Mexico (Guzmán et al., l 993a). This and their colleagues R. Valenzuela at ENCB, and allergy is produced by the large number of spores L. Guzmán-Dávalos at IBUG, helped with discus­ from the basidiomata. that sometimes form a mist sions, critica! revisions, herbarium studies, her­ of spores in r.he mushroom houses. Neverr.heless, barium loans, and/or bibliographic revisions. r.his adverse allergic reaction is not dangerous if Drs. O. K. Miller. R. T. Moore, D. N. Pegler, persons who are working inside r.he houses change R. H. Pe tersen, J. Royse, R. Vilgalys, S. P. Wasser, their activity for an external operation, for example, and J. Zervakis facilitated important biblio­ preparation of the substrate outside. The author graphic references, herbarium material. strains, himself observed r.hat when a person is insi_de a and/or opinions. This article was critically revised mushroom house with plenty of basidiomata in bv L. Guzmán-Dávalos. F. Ramfrez-Guillén, and - . ' production for a long period of time, headaches F. Tapia. ·. .,. and other symptoms such as cold or respiratory disorders are present. The symptoms change, how­ ever, according to the individuals' .sensitivities. REFERENCES Fatigue, mild headache. sinus pressure, coughing, mild difficulty in brearhing, pain in the limbs and Acosta-U rdapilleta L., Bautista N., López L.. Portugal D .. joints. and generalized malaise or ill feeling (in­ Montiel E.. and Mora V. \-I. 1995. Almacenamiento íluenzalike symproms). even fever of 39--J.OºC, de esperadas a baja ¡emperacura y su influencia sobre el which disappeared wir.hout trearment. are common desarrollo mi celial de cepas multiespóricas de P!euroms signs of the allergy produced by the spores of osrreatus . .Hic o/ Neorrop Apl. 8, 65-7 l. Acosta-Urdapilleta L., Bustos-Zagal G., and Portugal­ Pleurotus cultivated in commercial houses.' Portugal D. 1988. Aislamientos y caracterización de ·v. Recently, the author carne across the book by cepas de Pleurorns osrreatus y su cultivo en residuos agro­ Ying et al. ( 1987) where important inforrnation on industriales en d Estado de Morelos. Rev Me:c :vfico/ . ./., r.he medicinal properties of P. fe rulae, P ostreatus, 13- 20. P comucopiae. P spodoleucus, P. cirrinopileatus, Aguilar A., Martínez-Carrera D., Parra F., Sánchez­ and P. ulmarius is discussed. P. ferulae cures gas­ Hernández M., Morales P., and Soba! M. 1993. Anál­ isis económico y financiero de una planea rural de pro­ tropathy and kills insects and worrns. P ostreatus ducción de hongos comestibles (Píeurorus) en Cueczalan. cures lumbago arid~Rainful legs, numbed limbs, Puebla. México. Mico/ Neotrop Ap/. 6, 81- 94. and discomfort in tendons and blood vessels. The Anderson N.A .. Wang S. S., and Schwandt J. W. 1973. water extract is used against sarcoma and Ehrlich The Pleurotus osrreaws-sapidus species complex. Mvco!o­ carcinoma. P. comucopiae is used also against gia. 65, 28-35. Aslan-Azizi K., Shamala T. R., and Sreekantiah K. R. sarcoma and Ehrlich carcinoma. Aqueous extract 1990. Culcivacion of Pleurows sajor-cajll on certain agro­ of P. spodoleucus was tested against the growth of industrial wasres and ucilizacion of che residues for cellulase sarcoma. P. _citrinopileatus and P. ulmarius may and o-xylanase production. Mllshr J Tropics. 10. 21-26. cure pulmonary emphysema and P.' ulmarius is Bano Z., Bhagya S., and Srinivasan K. S. 1981. Essential also used against sarcoma and Ehrlich carcinoma. amino acid composition and proximate analysis of che mushrooms Pleurotus eous and P.jlorida. Mushr Newslett Tropics, 1 (3), 6-10. Bano Z. and Rajaraihnam S. 1982. P/e¡¡rotus mushroom as a nutritious food. [n: Tropical mushrooms. Biological ACKNOWLEDGMENTS nature and cultivation methods. Chang S. T. and Quimio T. H.. eds .. The Chinese Universiry Press. Hong Kong, The author is thankful for the help he has re­ pp. 363-380. Barron G. L. 1977. The nematode-desrroying fungi. Cana­ ceived from many colleagues in his srudies on dfan Biol Pubis, Gueiph (Onrario), 140 p. Pleurotus since 1970. Among them, Drs. Alexan­ Barron G. L. and Thorn R. G. 1987. Desrruction ofnema­ der H. Srnith. Albert Pilát. and Rolf Singer were todes by species of Pleurotlls. Can J Bot. 64, 774-778. of great help with opinions and also inforrnation Bas C. 1990. Pleuroraceae Over. ex Kühner. In: Flora from their laboratories. His co-wokers at Xalapa, Agaricina Neeriandica. vol. 2, Bas C. , Kuyper T. W..

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