全 文 ：Social and Biotechnological Studies of Wild Edible Mushrooms
in Mexico , with Emphasis in the Izta-Popo
and Zoquiapan National Parks
Perez-Moreno J .
?
, Martinez-Reyes M . , Lorenzana Fernandez A . ,
Carrasco Hernandez V . , Mendez-Neri M.
( Microbiología, Edafología, Colegio de Postgraduados, Km 36 .5 carr . México-Texcoco,
Montecillo, Texcoco, Estado de México, CP 56230 , MEXICO)
Abstract: In Mexico, there is an ancestral traditional knowledge related to wild edible mushrooms . Currently, more than
200 species of wildmushrooms are consumed in the country and morethan 100 are sold in traditional markets . In this pa-
per, first, the levels of commercialization of wild ediblemushrooms in Mexico are analyzed . Then, the species sold in the
area of influence of the Izta-Popo and Zoquiapan National Parks, located inCentral Mexico and their phenological patterns
are presented . Finally, an evaluationof the effect of pine growth under greenhouseconditions and field survival as a result
of inoculation with three ectomycorrhizal ediblefungi is presented . In theregion, more than 100 names areused to denote
edible species . The highest numbers of fungal specieswere recorded in July and August . Ground pileus have been used as
a successful source of ectomycorrhizal inoculum . Until now, the best results have been obtainedwith species of Hebeloma,
Laccaria and Suillus under greenhouseconditions . Evident increases in survival of plants inoculated withamixtureof these
species have been recorded under field conditions compared with non- inoculated plants .
Key words: Phenological patterns; Ectomycorrhizal fungi ; Inoculation; Pinus; Field survival
CLC number : S 646 Document Code : A Article ID: 0253 - 2700 ( 2009) Suppl.ⅩⅥ- 055 - 07
Introduction
Mexico holds large biological and cultural diversi-
ties . It has been considered a mega diverse country,
possessing, together with 12 other countries, 80% of
theplanet′s biodiversity and alone has around 30 000
plant species and 45 vegetation types ( Conabio,
1998) . Somegroupsof organisms, whose biological di-
versity in the country is huge, are especially important
worldwide . For example, the country is first in the
number of species of ectomycorrhizal ( ECM ) trees of
major forest importance, such as Pinus (with 72 taxa)
(Perry, 1991) and Quercuswith 150 species (Valencia,
2004) . As a result of this peculiar combination of great
biological and cultural diversity (includingmore than 50
ethnic groups) , there is profound traditional knowledge
of wild fungi . The use of wild edible mushrooms in
Mexico goes back to pre-Hispanic times, as evidenced
by linguistic and ethnological tests . In the country more
than 200 wild mushrooms are consumed (Villarreal and
Pérez-Moreno, 1989) , andmore 1000 common names in
thedifferent native languages are used to denote these
ediblemushrooms species (Guzmán, 1997) . Consider-
ing the abovescenario, a project that includes taxonom-
ic, social , ecological and biotechnological aspects of
wild ediblemushrooms has been carriedout currently in
the Izta-Popo and Zoquiapan National Parks and its sur-
rounding areas, inCentral Mexico, funded by theMexi-
can government ( through the project SEMARNAT-
CONACYT 2004 - 01 - 45) from2005 to 2008 . This pa-
per presents somesocial and biotechnological information
of thewild ediblemushroomsof Mexico, including: i) a
summary of the levelsof commercializationof wild edible
mushrooms in Mexico; ii ) a phenological assessment of
68 wild edible mushrooms in nine markets in Central
Mexico; and iii) an evaluationof theinfluenceof inocu-
lation with three edible ECM mushrooms on pinegrowth
under greenhouse conditions and survival of inoculated
plants under field conditions .
云 南 植 物 研 究 2009 , Suppl . ⅩⅥ : 55～61
Acta Botanica Yunnanica
? ?Author for correspondence; E-mail : jperezm@ colpos.mx??jepemo@yahoo. com. mx; Tel + 52 595 9520200 ext . 1280
Materials and Methods
Studied area The Izta Popo and Zoquiapan Na-
tional Parks are located inCentral Mexico, between 18°
59′and 19°16′25″N and 98°34′54″and 98°16′25″W .
The area is covered mainly with coniferous and broad-
leaf forests, including: i ) pine forests (mainly Pinus
hartwegii and P . pseudostrobus) from2 500 to 4 000 m
altitude; ii) fir forests ( Abies religiosa) from 2 500 to
3 500 maltitude; iii ) mixed forests includingecotones of
Pinus spp ., Alnus spp . and Quercus spp . from2 500
to 3 500 maltitude, and iv) subtropical moist montane
forests from 2 500 to 2 800 m altitude ( Izta Popo Na-
tional Park, 2007 ) . These kinds of forests favor the
development and diversity of wild edible mushrooms .
The park includes 22 municipalities and their area of
environmental and social influence affects three Mexi-
can States: Mexico, Puebla and Morelos, and also
Mexico City . First gatherers, hunters and fishers ar-
rived to the area, at least, 22 000 years ago as shown
by archaeological evidence . Collection of wild edible
mushrooms in the region is therefore ancestral anddefi-
nitely started fromPrehispanic times in nativegroupsof
náhuatl origin .
Phenological assessment To register the sea-
sonal variations of the species of wild ediblemushrooms
that are marketed in the area of influence of the Izta-
Popo Zoquiapan National Park, 50 visits were made to
9 markets, during 2006 . The sampled markets of this
area are located in the state of Mexico and Mexico
City . Identification of wild edible mushrooms was car-
ried out by using routine mycological techniques and
both macro and micromorphological characteristics of
themushrooms . Authorities of fungal species are those
recognized in Index Fungorum (2008) .
Biotechnological studies In the inoculation ex-
periments two pine species and three edible ECM
mushrooms were studied . Mushrooms were chosen
among species commonly sold in the local markets . In
thepreparation of ECM inoculum, pileus of Hebeloma
mesophaeum ( Pers .) Quél . and Laccaria bicolor
(Maire) P . D . Orton, and hymeniaof Suilluspseudo-
brevipesA . H . Sm & Thiers were used . These struc-
tures were dehydrated below 36℃ , then ground and
filtrated by a 1 mm mesh to obtain uniform particle
size . Seeds wereplanted inblack plastic140 cm3 tubes
that contained amixtureof sawdust-sand-soil in a 2∶2∶
1 proportion . Each plant was inoculated with at least
107 to 108 spores . Two inoculations were performed to
increase the probability of colonization . ECM inocu-
lums was placed in holes made on the surface of the
substrates contained in the tubes . The plants were irri-
gated with distilled water for 240 days when they are
necessary . Shoot and root dry weights were recorded
240 days after planting with plants that were previously
dried at 80℃ . Shoot and root total N and P of each
plant were calculated following Bremner ( 1965 ) and
Olsen et al . (1954) , respectively . Thepercentagesof
ECM colonizationwere calculated for all plants . Addit-
ionally, morphological characterization of the morpho-
types found was conducted ( Agerer, 1991 ) to assess
the presenceof themushrooms used as inoculums . The
data obtained for different variables were subjected to
an analysis of variance and the Tukey test of compari-
son of means (α= 0 .05) (SAS Institute, 1999) . Sur-
vival of inoculated and non-inoculated plants was eval-
uated under field conditions in a parallel set of Pinus
greggii plants kept under greenhouse conditions for 12
months . Evaluation of plant survival was carried out
92 , 112 , 170 and 262 days after seedling transplant-
ing . The selected site lacked forest during the last 30
years .
Results
Commercial wild edible mushrooms in Mexico
The use and commercialization of wild edible
mushrooms is very common in Central and Southern
Mexico and ingeneral terms, it can begrouped in four
categories:
1) Self-consumption: A largenumber of wild edi-
ble mushrooms have agreater value than exchange val-
ue and therefore they are self-consumed by native
mushroom collectors . This category includes: i ) spe-
cies with a short shelf life; ii) species with usual low
natural productions; and iii ) species highly valued by
local collectors for their special taste . Occasionally
some speciesbelonging to this category aresold inmar-
65 云 南 植 物 研 究 增刊ⅩⅥ
kets as well , for example when natural productions are
much more than usual . Around 100 species are
grouped in this category .
2) Direct commercialization by native collectors:
Frequently, mainly in the rural areas of the country,
native collectors directly sell mushrooms to consumers,
without brokers or middlebusinessmen . In these areas,
collectors usually travel very long distances on foot try-
ing to gather the highest possible amounts of mushro-
oms . Mushroom collection can start very early in the
morning, around 4: 00 a.m . and finish around 5: 00
p.m, this is more than 12 hours of work daily . In
these cases, there is a low-scale commercialization and
frequently each collector commercializes no more than
20 kilograms at atime . Thecommercialization is usual-
ly accomplished with mixtures of species in pileus rath-
er than in complete kilograms . In these kinds of mar-
kets, prices are not fixed because bargain is a very
common practice . Even when mushroomcollection is a
familiar activity, sellers in these kinds of markets are
usually women, maybe because of their great capabili-
ties to negotiate prices . More than 100 species are
commercialized in Mexican traditional markets .
3 ) Commercialization through intermediaries . In
large cities, such asMexico City, saleof wild mushro-
oms is mainly through intermediaries . Wild mushrooms
are transported from areas where they are bought from
nativegatherers to wholesalemarkets by vehicles . It is
very frequent that the chain is made up of three or four
intermediaries . Price bargaining is frequent that at each
change of hands . Sales in these largemarkets inMexico
City, for example, take place at different times . In
“Central de Abastos”market sale begins at 4: 00 a.m .
and ends around 7: 00 a.m ., while in“La Merced”
market it begins around 6: 00 a.m . and ends around 3:
00 p.m . Some of the large markets are truly regional
collection centers given the simultaneous commercializa-
tion of wild mushroomsfromdifferent parts of the coun-
try . Weekly quantities of commercialized wild mushro-
oms in these markets are measured in tons .
4) Exportation to international markets . A re-
duced number of ECM wild mushrooms species of Mex-
ico is exported to international markets, mainly to the
United States, several European countries and Japan .
Among these, Amanita caesarea s . l ., Boletus edulis
s . l ., Morchella spp ., Cantharellus cibarius and Tri-
choloma matsutake are the principal species . Although
there are admirable attempts, for example of the Mixe
group of Oaxaca, the legal framework in general and
thedegreeof social organization for wild ediblemushro-
oms export fromMexico is quite incipient, as opposite
to other countries with also great ancestral mycological
tradition, like China . Frequently, most of the profits
remain in the groups of intermediaries who buy com-
mercially valuable wild edible mushrooms from local
gatherers at very low prices and at the same time de-
mand very high quality . The prices paid to the local
gatherers vary conspicuously depending on the region
and the degree of information to which the gatherers
have . In general , prices paid to native collectors of
mushrooms subject to exportation, aresadly established
in theblack market . Dozensof Tons of wildmushrooms
are exported yearly from different parts of Mexico but
becauseof the lack of strict legal regulation, exact data
on the commercializationof the different species arenot
accurately known . However, the exportation of these
wild edible mushrooms from Mexico, definitely has a
commercial value, withgreat social importance for poor
local communities wherecollectionof wild ediblemush-
rooms is usually carried out in every year .
Phenology of wild edible mushrooms
Three phenological patterns ( Table 1 ) , of those
proposed for wild edible mushrooms of Mexico, were
identified: i ) 44% of the species presented a short
early fructification pattern (May to August) ; ii ) 31%
of the species presented an early prolonged fructifica-
tion pattern ( July to December) ; and iii ) 25% of the
species presented a short fructifications in themiddleof
the season pattern ( J uly to October) . Most of the spe-
cies were recorded in the middle of the rainy season,
fromJuly toAugust (Table1 ) . In thestudied area, we
recorded more than 100 common names used to denote
wild edible mushrooms . However, in general terms, a
conspicuous loss of mycological traditional knowledge is
occurring in the area in part originated by: i ) agricul-
tural crisis; ii ) influenceof massive communicationme-
75增刊ⅩⅥ Perez-Moreno J . et al . : Social and Biotechnological Studies of Wild Edible Mushrooms in . . .
dia; and iii ) modification of familiar and social values .
However, according to our observations there are some
areas in the region, mainly in small towns, wherenative
people still retain its traditional culture and subsistence
pre-Hispanic practices, including collection and con-
sumption of wild edible mushrooms .
Table 1 Phenological patterns of wild edible mushrooms commercialized in nine the markets in Central Mexico in 2006 *
Pp Species
Trophic
group
Months
M J J A S O N D
1 ?? 2 f1 ?2 1 R2 1 2 >1 2 1 *2 1 r2 1
a Agaricus campestris Fr . S + + +
a Amanita vaginata var -. vaginata ( Bull .) Lam . ECM + + + +
a Armillaria mellea ( Vahl) P . Kumm . S + + + + +
a Boletus clavipes ( Peck) Pilát & Dermek ECM + + + +
a B ?. luridus Schaeff . ECM +
a Retiboletus griseus ( Frost) Manfr . Binder & Bresinsky ECM + +
a Clavulina coralloides (L .) J . Schr?t . ECM +
a Entoloma clypeatum (L .) P . Kumm . ECM + +
a Flammulina velutipes ( Curtis) Singer S +
a Gomphus floccosus ( Schwein .) Singer ECM + +
a Hebeloma alpinum ( J . Favre) Bruchet ECM + + + + +
a H ?. leucosarx P . D . Orton ECM + + + + +
a H ?. mesophaeumvar. mesophaeum ( Pers .) Quél . ECM + + + + +
a Hygrophoropsis aurantiaca ( Wulfen) Maire S + + + + +
a Hygrophorus hypothejus ( Fr .) Fr . ECM + + +
a Hypomyces lactifluorum ( Schwein .) Tul . & C . Tul . P + + + + +
a H ?. macrosporus Seaver P + +
a LycoperdumpyriformeSchaeff . S +
a Pluteuscervinus var ?. cervinus (Schaeff . ex: Fr .) Kum . S + +
a Ramaria stricta var ?. concolor Corner S + + + + + +
a R ?. cystidiophora ( Kauffman) Corner ECM ? + +
a R ?. fennica var N. fennica ( P . Karst .) Ricken ECM ? + +
a R ?. rubiginosa Marr & D. E . Stuntz ECM ? + + + +
a Ramaria rubrievanescens Marr & Stuntz ECM ? + + + +
a R ?. rubripermanens Marr & D ?. E . Stuntz ECM ? + + + +
a R ?. sanguinea ( Pers .) Quél . ECM ? + + + +
a R ?. stricta ( Pers .) Quél . S + +
a Russula brevipes Peck ECM + +
a R ?. olivacea ( Schaeff .) Fr . ECM + + + +
a Suillus pseudobrevipes A . H . Sm . & Thiers ECM + +
b Gyromitra infula ( Schaeff .) Quél . ECM + + +
b Hygrophorus russula ( Schaeff .) Kauffman ECM + + +
b Amanita crocea ( Quél .) Singer ECM +
b Ustilago maydis (C . D .) Corda P + + + + + + +
b Clavulina cinerea f ?. cinerea (Bull .) J . Schr?t . ECM + + +
b
C ?. cinerea f ?. sublilascens (Bourdot & Galzin) Bon
& Courtec .
ECM + + + + + +
b
Ampulloclitocybe clavipes ( Pers .) Redhead,
Lutzoni , Moncalvo & Vilgalys
ECM + + + +
b Helvella elastica Bull . ECM + + + +
b Pseudocraterellus undulatus ( Pers . Rauschert) ECM + + + + +
b Laccaria proximella Singer ECM + + + +
b Lycoperdon perlatumPers . ECM + + + +
B Pleurotus cornucopiae ( Paulet) Rolland S + + +
B Ramaria aff ?. gelatinosa Holmsk . ECM ? + + + +
B R ?. holorubella ( G. F . Atk .) Corner ECM ? + + + + + +
B R ?. pallida ( Schaeff .) Ricken ECM ? + +
B R ?. rasilispora Marr & D. E . Stuntz ECM ? + + + + + +
B Russula mexicana Burl . ECM + + +
c Amanita caesarea (Scop .) Pers . ECM + + + + + + + +
c A ?. franchetii (Boud .) Fayod ECM + + + + + + + + +
85 云 南 植 物 研 究 增刊ⅩⅥ
Continue table 1
Pp Species
Trophic
group
Months
M J J A S O N D
1 ?? 2 f1 ?2 1 R2 1 2 >1 2 1 *2 1 r2 1
c A ?. fulva ( Schaeff .) Fr . ECM + + + + + + + +
c A ?. rubescens var. rubescens Pers . ECM + + + + + + + + +
c Boletus edulis Bull . ECM + + + + + + + +
c B ?. pinophilus Pilát & Dermek . ECM + + + + + + + + +
c Cantharellus cibarius var ?. cibarius Fr . ECM + + + + + + + + + + + + +
c Clitocybegibba ( Pers .) P . Kumm . ECM + + + + + + + +
c Gymnopus dryophilus ( Bull .) Murrill S + + + + +
c Helvella lacunosa Afzel . ECM + + + + + + + +
c Laccaria amethystina Cooke ECM + + + + + + + + +
c L ?. bicolor (Maire) P ?. D . Orton ECM + + + + + + + + + + +
c L ?. laccata ( Scop .) Cooke ECM + + + + + + + + + + + + + +
c L ?. proxima ( Boud .) Pat . ECM + + + + + + + + + + + + +
c Lactarius deliciosus (L .) Gray ECM + + + + + + + +
c L ?. indigo (Schwein .) Fr . ECM + + + + +
c L ?. salmonicolor R . Heim& Leclair ECM + + + + + + + +
c Lyophyllumdecastes ( Fr .) Singer ECM + + + + + + + + + +
c Morchella esculenta (L .) Pers . ECM + + + + + + + + + + + +
c M ?. elata Fr . ECM + + + + + + + + + + +
c Russula delica Fr . ECM + + + + + + + +
* The studied marketswere: i) in Mexico city:“La Merced”,“ Jamaica”and“Central deAbastos”; ii) in the stateof Mexico:“Texcoco”,“Ozumba”,
“Amecameca”,“Chalco”,“San Rafael”and“Colonia?vila Camacho”. Pp= Phenological patterns: a= species with short early fructification ( May to
August) ; b= species with short fructification in the middle of the season ( July to October) ; c = species with early prolonged fructification ( J uly-August-
December) . Trophic group: S = saprobe; ECM = ectomycorrhizal ; P = parasite . 1 = first fortnight of the month; 2 = second fortnight of the month
Biotechnological essays
Ingeneral terms, plant inoculationwith any of the
three evaluated ECM ediblemushrooms produced a sig-
nificant increase in terms of dry weight of the shoot and
roots compared with non-inoculated plants . A similar
trend was observed in the case of N and P contents .
The highest beneficial effect was produced in the case
of plants inoculated with Hebeloma mesophaeum ( Table
2) . Therewas no evidence of synergistic or competitive
effect when simultaneous inoculation with thethreeECM
fungi was carried out . The percentages of total ECM
colonization were 58, 57 , 76 and 59% for P . greggii
and 84 , 77 , 89 and 63% for P . pseudostrobus, in pla-
nts inoculated with H. mesophaeum, L . bicolor,
S. pseudobrevipes and the mixture of the three ECM
species, respectively . Meanwhile, ECM colonization in
non-inoculated plants was 19 and 22% for P . greggii
and P . pseudostrobus, respectively . The morphotype
analysis revealed that ECM colonization according to
the inoculated species ranged from 15 to 69% in
P . greggii and from 41 to 89% in P . pseudostrobus .
Despite the fact that seedling transplantingwasmade at
the end of the rainy season, the percentage of survival
was up to 100% , for all treatments 92 and 112 days
after transplanting . In contrast 170 and 262 days after
transplanting, during dry season, plant survival de-
creased distinctly compared with the inoculated edible
ECM fungi . In general terms, thehighest percentageof
survival was observed in plants inoculated with the
combination of threeECM fungi . Plants inoculatedwith
either S. pseudobrevipes or L . bicolor presented higher
survival percentages than those inoculated with H.
mesophaeumor non-inoculated (Table 3) . It has been
considered that someECM fungi can confer drought tol-
erance to their associated hosts . In the caseof S. pseu-
dobrevipes such probable tolerance can be induced by
the presence of abundant rhizomorphs which can dra-
matically, increase the absorption area ( Duddridge et
al. , 1980) .
Discussion
Increments in plant biomass had been documented
as a result of inoculation with genera of some of the
ECM mushrooms studied in our work . For example,
95增刊ⅩⅥ Perez-Moreno J . et al . : Social and Biotechnological Studies of Wild Edible Mushrooms in . . .
Table 2 Dry weight and nutrient contents of two pine species inoculated or no with three edible
ectomycorrhizal mushrooms, 240 days after sowing under greenhouse conditions
Variable and
treatments
Plant species and parts of the plant
Pinus greggii Pinus pseudostrobus
Shoot Root Total Shoot Root Total
Dry weight
Nip 314 ?. 9 c 190 !. 5 d 505 _. 4 c 291 .5 c 137 #. 3 c 428 a. 8 b
piHm 770 ?. 8 a 276 !. 2 b 1 ?052 .1 a 540 .5 a 218 #. 5 a 759 a. 0 a
piLb 592 ?. 5 b 259 !. 8 bc 852 _. 3 b 413 .7 ab 160 #. 9 b 574 a. 7 ab
piSp 612 ?. 8 b 248 !. 6 cd 861 _. 8 b 389 .5 b 203 #. 6 a 618 a. 1 ab
pi LHS 735 ?. 3 a 296 !. 6 a 1 ?026 .8 a 490 .6 a 234 #. 0 a 724 a. 6 a
N content
sNip 2 ?. 4 c 1 . 7 c 4 ?. 1 c 3 . 0 c 0 .9 c 3 .9 c
piHm 9 ?. 5 a 2 . 7 b 12 :. 2 a 4 . 6 b 1 .8 b 6 ?. 4 bc
piLb 9 ?. 4 a 2 . 8 b 12 :. 2 a 6 . 4 a 2 .1 ab 8 ?. 5 a
piSp 5 ?. 8 b 2 . 0 c 7 ?. 8 b 5 . 0 b 1 .8 b 6 .8 b
pi LHS 8 ?. 8 a 3 . 4 a 12 :. 2 a 5 . 8 a 2 .5 a 8 ?. 3 a
P contents
Nip 0 ?. 31 b 0 . 27 b 0 ?. 58 d 0 . 25 c 0 .13 c 0 .38 c
piHm 0 ?. 93 a 0 . 47 a 1 ?. 41 a 0 . 63 a 0 .31 ab 0 .94 a
piLb 0 ?. 70 a 0 . 40 a 1 ?. 11 b 0 . 65 a 0 .30 ab 0 .95 a
piSp 0 ?. 63 a 0 . 19 c 0 ?. 82 c 0 . 45 b 0 .23 b 0 .68 b
pi LHS 0 ?. 72 a 0 . 39 a 1 ?. 11 b 0 . 60 ab 0 .38 a 0 .98 a
Note: Nip= non- inoculated plants; piHm= plants inoculated with Hebeloma mesophaeum; piLb = plants inoculatedwith Laccaria bicolor; piSp= plants
inoculated with Suillus pseudobrevipes; piLHS = plants inoculated with L . bicolor, H . mesophaeumand S. pseudobrevipes . Numbers with different letters in
each part of the plant, in each variable, are different according to Tukey test ( P = 0 . 05) . n= 20 .
Table 3 Percentage of survival of Pinus greggii plants inoculated with three edible ectomycorrhizal fungi and
non- inoculated plants, during 9 months after transplanting
Treatments
Days after transplanting
92 112 ?170 Y262
Non- inoculated plants 100 9100 ?44 G19
Plants inoculated with Hebeloma mesophaeum 100 9100 ?90 G10
Plants inoculated with Laccaria bicolor 100 9100 ?100 Y50
Plants inoculated with Suillus pseudobrevipes 100 9100 ?93 G57
Plants inoculated with LHS * 100 9100 ?100 Y100
* a mixtureof with L . bicolor, H. mesophaeum and S. pseudobrevipes . n= 16
Sudhakara-Reddy and Natarajan ( 1997 ) reported in-
crements of 218% and 182% in shoot and root dry
weight, respectively, as a result of inoculation with
Laccaria laccata, relative to non-inoculated Pinuspat-
ula plants, twelvemonths after inoculation in sterilized
soil . Additionally, several studies have found that si-
multaneous inoculation with different ECM mushrooms
can produce larger increments in biomass in inoculated
plants than inoculation with a single species ( Parladé
and?lvarez, 1993; Sudhakara-Reddy and Natarajan,
1997) . Sudhakara-Reddy and Natarajan (1997) inocu-
lated Pinus patula with Laccaria laccata and Thele-
phora terrestris and found a synergetic effect in shoot
dry weight as a result of simultaneous inoculation with
these species . However, in our study simultaneous in-
oculation with the three ECM species studied did not
produce a synergistic effect, even though a generally
beneficial effect was maintained . Several authors have
reported that inoculation with ECM mushrooms causes
an increment in the total nutrient content of both tem-
perate and tropical trees and bushes, mainly that of N
and P ( Turjaman et al. , 2006; Nara, 2006; Tibbett
and Sanders, 2002) . In our case we used early-stage
ECM species: Laccaria, Hebeloma and Suillus (Ma-
son et al. , 1983) . Usually early or intermediate stage
ECM species (asopposed to late-stage species) require
only small amounts of carbon from their hosts and low
concentrations of nitrogen and phosphorus (Gibson and
Deacon, 1990; Bergemann and Miller, 2002 ) . These
species usually produce increases in plant biomass in
06 云 南 植 物 研 究 增刊ⅩⅥ
the greenhouse or nursery probably because of their
preference for mineral sources of N and P ( Rincón et
al. , 2001; Sudhakara-Reddy and Natarajan, 1997;
Chu-Chou and Grace, 1985 ) . It was reported that
ECM symbiosis can cost7 to 30% of theC fixedby the
plants ( Leake et al. , 2004 ) . Although this cost may
be considered high, once themycelial networks are es-
tablished, there is the possibility of substantial translo-
cation of N and P ( Pérez-Moreno and Read, 2000 ,
2001a, b) ; thus, ECM symbiosis in general terms is
considered mutuality ( Read and Pérez-Moreno,
2003) . However, our inoculation experiments showed
that groundpileus can beused as asuccessful sourceof
ECM inoculums for pines . Studies on the other species
of ECM edible fungi and plants would be highly desir-
able .
Acknowledgment: Financial support is acknowledged from
project SEMARNAT-CONACyT-2004 - 01 - 52:“Los hongos sil-
vestres comestibles del Parque Nacional Izta-Popo Zoquiapan y
Anexos”.
References:
Ager ?er R , 1991 . Characterization of ectomycorrhiza [ J ] . MethodsMicro-
biol , 23 : 25—73
Berg ?emann SE, Miller SL , 2002 . Size, distribution, and persistence of
genets in local populations of the late-stage ectomycorrhizal basidio-
mycete, Russula brevipes [ J ] . New Phytologist, 156: 313—320
Brem /ner JM, 1965 . Total nitrogen [ A ] . In: Black CA (ed) Methods of
soil analysis Part 2 , Agronomy 9 [ M] .Madison WI : AmericanSoci-
ety of Agronomy, 1149—1178
Chu- %Chou M , Grace LJ , 1985 . Comparative efficiency of the mycorrhizal
fungi Laccaria laccata, Hebeloma crustuliniforme and Rhizopogon
species on growth of radiate pine seedlings [ J ] . New Zealand J our-
nal of Botany, 23 : 417—424
Cona bio, 1998 . Ladiversidad biológica de México: estudio de un país .
México: Comisión Nacional para el Conocimiento y Uso de la Biodi-
versidad (CONABIO)
Dudd 3ridge JA , Malibari A , Read DJ , 1980 . Structure and function of
mycorrhizal rhizomorphs with special reference to their role in water
transport [ J ] . Nature, 287: 834—836
Gibs ?on F , Deacon JW, 1990 . Establishment of ectomycorrhizas in aseptic
culture: Effects of glucose, nitrogen and phosphorus in relation to
successions [ J ] . Mycol Res, 94 : 166—172
Guzm :án G, 1997 . Los nombres de los hongos y lo relacionado con ellos
en América Latina . Xalapa, México: Instituto de Ecología
Inde ?x Fungorum, 2008 . www. indexfungorum.org [ OL ]
Izta ?-Popo National Park, 2007 . www. iztapopo. conanp.gob. mx? [OL]
Leak ?e J , Johnson D, Donnelly D et al. , 2004 . Networks of power and
influence: the role of mycorrhizal mycelium in controlling plant com-
munities and agroecosystem functioning [ J ] . Canadian J ournal of
Botany, 82 : 1016—1045
Maso ?n PA , Wilson J , Last FT, 1983 . The concept of succession in rela-
tion to spread of sheathing mycorrhizal fungi on inoculated tree seed-
lings growing in unsterile soils [ J ] . Plant and Soil , 71 : 247—256
Nara ?K , 2006 . Ectomycorrhizal networks and seedling establishment dur-
ing early primary succession [ J ] . New Phytologist, 169: 169—178
Olse ?nSR , Cole CV, Cantable FS et al. , 1954 . Estimation of available
phosphorus in soils by extraction with sodium bicarbonate, circular
939 . Washington, DC: US Deparment of Agriculture
Parl ?adé J , ?lvarez IF , 1993 . Coinoculation of aseptically grown Douglas
fir with pairs of ectomycorrhizal fungi [ J ] . Mycorrhiza, 3 : 93—96
Pére ?z-Moreno J , Read DJ , 2000 . Mobilization and transfer of nutrients
from litter to tree seedlings via thevegetative myceliumof ectomycor-
rhizal plant [ J ] . New Phytolologist, 145: 301—309
Pére ?z-Moreno J , Read DJ , 2001a . Exploitation of pollen by mycorrhizal
mycelial systems with special reference to nutrient recycling in boreal
forests [ J ] . Proceedings of the Royal Society of London, 268:
1329—1335
Pére ?z-Moreno J , Read DJ , 2001b . Nutrient transfer from soil nematodes
to plants: A direct pathway provided by themycorrhizal mycelial net-
work [ J ] . Plant Cell Environment, 24 : 1219—1226
Perr ?y J , 1991 . Thepinesof Mexico and Central America [M ] . Portland,
Oregon: Timber Press
ReadDJ , Pérez-Moreno J , 2003 . Mycorrhizas and nutrient cycling in eco-
systems - a journey towards relevance [ J ] . New Phytologist, 157:
475—492
Rinc ?ón A, ?lvarez IF , Pera J , 2001 . Inoculation of containerized Pinus
pinea L . seedlings with seven ectomycorrhizal fungi [ J ] . Mycorrhi-
za, 11 : 265—271
SAS ?Institute, 1999 . SAS User′s Guide, ver . 8 . 0 . Cary, NC: SAS In-
stitute Inc
Sudh ?akara-Reddy M , Natarajan K , 1997 . Coinoculation efficacy of ecto-
mycorrhizal fungi on Pinus patula seedlings in a nursery [ J ] . My-
corrhiza, 7 : 133—138
Tibb ?ett M , Sanders FE, 2002 . Ectomycorrhizal symbiosis can enhance
plan nutrition trough improved access to discrete organic nutrient
patches of high resource quality [ J ] . Annals of Botany, 89 : 783—
789
Turj ?aman M, Tamai Y , Segah H et al. , 2006 . Increase in early growth
and nutrient uptake of Zorrea seminis seedlings inoculated with two
ectomycorrhizal fungi [ J ] . Journal of Tropical Forest Science, 18 :
243—249
Vale ?ncia AS, 2004 . Diversidad del género Quercus ( Fagaceae) en
México [ J ] . Boletín de la Sociedad Botánica deMéxico, 75 : 33—53
Vill ?arreal L , Pérez-Moreno J , 1989 . Los hongos comestibles silvestres de
México, un enfoque integral [ J ] . Micología Neotropical Aplicada, 2 :
77—114
16增刊ⅩⅥ Perez-Moreno J . et al . : Social and Biotechnological Studies of Wild Edible Mushrooms in . . .