全 文 ：Achievements and Challenges of Rrsearch on Truffles in China
?
WANG Yun
1 , 2 * *
, LIU Pei-Gui
1
??
( 1 Key Laboratory of Biodiversity and Biogeography, Kunming Instituteof Botany, Chinese Academy of Sciences,
Kunming 650204 , China; 2 New Zealand Institute for Plant & Food Research Ltd,
Private Bag 4707 , Christchurch 8104 , New Zealand)
Abstract: Research on truffles has made agood progress in China since the last two decades . Before 1989 only 8 Tuber
specieswere recorded . Since then, 25 species in Tuber and 1 in Paradoxa have been identified in China, of which are
morethan 50% new to science . More than 50 desert truffle species have also been recorded . These results suggest that
China has a rich truffleflora and might be one of centres of truffle diversification and distribution . Chinese black truffles,
the T. indicumcomplex, T. aestivum and T. pseudoexcavatum have been collected for consumption and trade locally for
many years . But theywerenot scientificallydescribed in Chinauntil recently . While Chinese black truffles theywerevirtu-
ally unknown internationally until theearly 1990swhen exports to international markets commenced . This has arousedgreat
concern and interest in the study of Chinese black truffles . Research results indicated that T. indicum is the predominate
truffle species exported to the international markets and a small amount of T. pseudoexcavatummixed with them . Genetic
and morphological variations of Chinese black truffle species are huge . T. indicum, T. sinense and T. himalayensearetoo
closely related to each other that they are indistinguishablemorphologically . The three species might be better called“the
T. indicum complex”. Harvesting and trading in Chinese black truffles is a multi-million dollar industry that has created
considerable incomefor small rural communities and farmers . Unfortunately, natural forests and plantations in whichtruffles
grow naturally have been damaged so much that created a big concern about their resources in danger . Protection of the
precious resources is urgent ! The first truffle plantation was established in Taiwan in 1989 and produced truffles in 1996 .
Other truffle plantations have been established in Hunan, Guizhou, Sichuan and Yunnan Province recently . It is said the
first group of truffle ascocarpswereproduced at the Guizhou plantation in2008 , but theother plantations havenot produced
truffles yet . The productionof truffle-infectedtrees andestablishment of truffleplantations areunderway and haveconsider-
able potential in China .
Key words: China; Truffles; Research; Cultivation
CLC number : S 646 Document Code: A Article ID: 0253 - 2700 (2009) Suppl .ⅩⅥ - 001 - 09
Diversity of Chinese truffle species
Research on truffles in last 20 years reveals that
China has big diversity of truffles . However, Chinese
ancient books had no records on truffle ( Tuber ) spe-
cies . The first Chinese truffle species, Tuber
taiyuanense to be recorded in China was in 1985 which
was described by Prof . B . Liu . Since then, research
on truffle diversity has expanded rapidly . Until now,
27 species and 1 variety have been recorded in China,
of which 26 species and 1 variety in Tuber and 1 in
Paradoxa . Among themthe 23 Tuber species and 1 va-
riety are“non-black coloured”and 3 are“Black co-
loured”truffle species ( known as“Chinese back truf-
fles”) , 12 of those species are new to science, and 7
are similar but not identical to known species and so
could be new species or subspecies ( Wang, 1990;
Zhang, 1990; Ren, 2003; Song, 2005; Wang et al. ,
2006a; Chen, 2007; Chen and Liu, 2007; Chen et
al. , 2008; Wang et al. , 2008 ) . A new combinationof
Parodoxa gigantospora (Y . Wang & Z. P . Li ) Wang
was published in 2008 based on Tuber species, Tuber
gigantosporumY . Wang & Z. P . Li ( 1991 ) because
云 南 植 物 研 究 2009 , Suppl . ⅩⅥ : 1～9
Acta Botanica Yunnanica
?
?? ?Author for correspondence; E-mail : wangy@crop. cri. nz; pgliu@ mail .kib. ac. cn
Foun ?dation items: This study was financially supported by the Joint Founds of theNational Science Foundation of China andYunnan ProvinceGov-
ernment (No . U0836604 ) , and Key Laboratory of Biodiversity and Biogeography, Kunming Instituteof Botany, ChineseAcademy of Sciences
( No . 0806361121) as well as Yunnan Program of Innovation to strong provinces by Science & Technology ( No . 2009AC013 )
its asci invariably contain only 1 spore . The first time
of the genus of Paradoxa has been found in China and
only the 2nd time that the genus has been found since
1935 when it was described by Mattirolo in Italy .
1 .“Non-black coloured”:
Tuber asa ?
Tuber borchii ?
Tuber borchii var. sphaerospermum
Tuber californicum
Tuber chenggonense #
Tuber dryophilum ?
Tuber aestivum
Tuber excavatum
Tuber furfuraceum *
Tuber huidongense *
Tuber leptoperidium #
Tuber liaotongense *
Tuber liui *
Tuber lyonii = T. texense
Tuber maculatum ?
Tuber latisporum *
Tuber pseudorufum #
Tuber puberulum ?
Tuber rapaeodorum ?
Tuber rufum ?
Tuber taiyuanense *
Tuber umbilicatum *
Tuber zhongdianense *
2 .“Black coloured”
The Tuber indicum complex:
Tuber indicum
Tuber sinense
Tuber himalayensis
Tuber pseudohimalayense
= Tuber seudoexcavatum
Tuber formosanum ( nom . invalid)
3 . Parodoxagigantospora
* New species, ? nov . sp . or subsp ., # not pub-
lished sp . nov .
Chinese black truffles which are locally called as
“wu-niang-tong” ( fruiting body without mother plant)
and“ song-mao-fuling” ( pine needle’ fungus) have
been collected for consumption and trade for many
years in Sichuan and Yunnan ( Zhang and Wang,
1990) . However, theywerenot scientifically described
in Chinauntil 1989 when theywerenamed as new spe-
cies of Tuber sinense by Tao et al . In fact, Tuber
sinense is almost identical to T. indicumwhich was de-
scribed in Asian (Cooke and Massee, 1892) based on
a truffle collection foundnear Mossorie in the northwest
Himalayas of India . Ninety seven years later Zhang &
Minter named another new species, T. himalayense
basedon a part of the same collection . In 1989 small
amount of Chineseblack truffleswas exported toGerma-
ny for appraisal . Since then, increasing quantities were
exported to international markets ( Yamanaka et al. ,
2001; Wang et al. , 2008) , which has created concern
and also interest in the study of these Chinese Tuber
species ( Fourrépers . comm, 1995; Wang and Hall ,
2001) . Finally, it has been found that T . indicum is
thepredominate truffle species exported to the interna-
tional markets and a small amount of T. pseu-
doexcavatum ( Wang et al. , 1998 ) mixed with them .
Last 16 years studies have revealed that genetic and
morphological variations of Chineseblack trufflespecies
arehuge . T . indicum, T. sinense and T. himalayense
are too closely related to eachother that they are indis-
tinguishable morphologically (Paolociioni et al. , 1997;
Mabru et al. , 2001; Song, 1995; Zhang et al. , 2005;
Wang et al. , 2006a; Chen, 2007) . Therefore, it would
be better to call them as“ the T. indicum complex”
(Wang et al. , 2006a, b; Wang et al. , 2008) . T. pse-
udohimalayense is a co-species of T. pseudoexcavatum
(Chen, 2007) . T. formosanum( nom . invalid) wasde-
scribed as a new species by Hu (1992) based on a col-
lection fromTaiwan but is, in fact, not avalid species .
Small amount of T. aestivum ( = T. uncinatum) is pro-
duced andsoldwith T. indicumlocally in Sichuan Prov-
ince (Fig.1 : a, b) . It is only recently that T. aestivum
( = T. uncinatum) was distinguished from T. indicum,
described scientifically and confirmed to be identical to
theEuropean collections by morphological and molecu-
lar methods (Song, 1995; Chen et al. , 1995) .
Additionally, over 50 non-tubers truffle species
belonging to 28 genera, 15 families havebeen found in
China in the last 20 years ( Tao, 1988; Wang & Liu
unpublished data) .
2 云 南 植 物 研 究 增刊ⅩⅥ
Fig. 1 Ascocarps ( a) and vertical section ( b) of Tuber aestivum
ZYGOMYCETES BASIDIOMYCETES
- Endogne - Alpova
ASCOMYCETES - Gautieria
- Elaphomyces - Gelopellis
- Balsamia - Gymnomyces
- Choiromyces - Hydnangium
- Fischerula - Hymenogaster
- Genebea - Hysterangium
- Genea - Melanogaster
- Geopora - Octavianiana
- Hydnobolites - Phlebogaster
- Hydnotrya - Rhizopogon
- Pachphloeus - Richoniella
- Leucangium( Picoa) - Sclerogaster
- Terfezia - Zelleromyces
Apparently, more research on non-tubers truffle
species will be needed in China (Fig. 2 : a, b)
Distribution of Chinese truffles
Only some part of Yunnan and Sichuan province
and small areas in the other parts of China have been
searched for truffles and so truffle distribution in China
has been found to bepredominately in south-west China
and very spotty in the restof China (Song, 2005; Chen,
2007; Chen and Liu, 2007; Wang et al. , 2008 ) . The
Tuber indicum complex, T. pseudoexcavatum, T. aes-
tivum, T. excavatum, T. huidongense, T. zhongdianese,
T. borchii var. sphaerospermum, Tuber liui , Tuber umbil-
icatum, Tuber latisporum and Parodoxa gigantospora
grow in south-west China (Yunnan and Sichuan) and
Taiwan . The rest of truffles have been found in both
south (Yunnan, Sichuan, Tibet, Hubei and Taiwan)
and north China ( Jilin, Liaoning, Inner Mongolia,
Beijing, Hebei , Shanxi , Gansu and Xinjiang) . It is
not surprised more truffle species and new distribution
region will bediscovered in China in the future .
Natural truffle habitats
In northern China truffle species have been found
in natural temperate forests and plantations, such as in
coniferous forestsor plantations of Pinus, Larix, Abies
and Picea and also in deciduous broad-leaved forestsof
Fig. 2 A fruit-body (a) and a vertical section ( b) of the species Geospora cooperi
3增刊ⅩⅥ WANG andLIU: Achievements and Challenges of Rrsearch on Truffles in China
Quercus, Carpinus, Tilia and Betula, as well as in
mixed forestsof pine and deciduousbroad-leaved trees .
For example, truffles have been found in mixed forests
of Pinus koraiensis with oak trees, plantations of P . -
koraiensis and Larix gmelinii in north east China;
mixed forestsof P . tabulaeformiswith Quercus mongol-
ica, Q. mongolica var. liangtungensis etc . and forests
and plantations of Picea spp . in north China; forests
and plantationsof Picea schrenkiana and Picea crassifo-
lia in northwest China (Fig.3) . Soils areusually acid .
In southern China, truffle species are found in
subtropical evergreen broad-leaved forests of Litho-
carpus, Castanopsis and Quercus, their second growth
of coniferous forests and plantationsof Pinus, Ketelee-
ria and Tsuga, or in deciduous broad-leaved forests of
Quercus, Castanea, Carpinus etc, and in mixed fore-
sts of pine and broad-leaved trees . Soils are usually
calcareous or sometimes acid soils . Rainfall ranges
from 300 to 600 mm annually .
The T. indicum complex, T. aestivum and
T. pseudoexcavatumshare thesamehabitats . They pro-
duced predominately from coniferous forests of Pinus
yunnanensis, P . armadii and Keteleeria evelyniana,
and their plantations (20 - 40 years old) . The conifer-
ous forests are the second growth of evergreen broad-
leaved forests of Castanopsis, Lithocarpus, Cyclobal-
anopsis and Quercus (1600 - 2600 m) . Chinese black
truffles also grow in evergreen broad-leaved forests but
production is much less . ( Fig. 4 ) . Coriaria nepalen-
sis, Pyracantha fortuneana, Phyllanthus emblica and
Ficus tikoua are plants commonly found in Chinese
black truffles producing areas whichmay be the indica-
tors of the habitats . Alnus ferdinandi-coburgii is often
found in theChineseblack truffles habitatsunder which
productionof ascocarps is higher than without themdue
to the soils withmoremoisten and N content . Soils are
normally calcareous with a pH of 7 .2 (5 .3 - 7 .9 ) and
have a high content of Ca, Mg and Fe, and high ratio
of Carbon to N ( Chen, 2007 ) . Compared with the
soils where T. melanosporum inhabits the calcareous
soils in south west China have lower pH due to heavily
leaching and coniferous needleswhichbringorganic ac-
ids to the soils . Chinese black truffles also inhabit in
moderately acid to acid red soils which developed from
lime stones or purple soli developed from purple sandy
stones (Tang et al. , 2005; Wang unpublished data) .
Tang et al. (2005) did intensive investigationonhabi-
tats of Chineseblack truffles in Pazhihua, Sichuan and
discovered that Chinese black truffles preferred slightly
acid oils with pH 6 .0 - 6 .5 on moderate slopes (20 -
30°) facing south . Though Chinese black truffles can
inhabit in the acid soils but webelieve they growbetter
and more productive in calcareous soils . García-Monte-
ro et al. ( 2008 ) and Geng et al. (2009) showed that
T. indicum and T. pseudoexcavatummycorrhizae grow
well in calcareous substrates rich in active carbonate .
Annual rainfall ranges from600 to 1000 mmwith more
than 2200 annual sunny hours .
Chinese black truffles produce distinctive br?lés
as T. melanosporum does, inside which grass, herba-
ceous plants and small shrubs died (Fig.5 ) . Ascocarps
form at depth of soils of 0 - 30 cm .
Chinese truffles and small mammals
Truffles producefruit-bodies belowground and re-
ly on animals, especially rodent, for dispersal of their
spores ( Johnson, 1996 ) . When small mammals eat
these truffles and deposit their spores elsewhere in their
feces . Therefore, mammals play an important role in
the maintenance of truffle mycorrhizal symbiosis and
abundance in truffle forest ecosystems . Some small
mammals opportunistically consume truffles and others
consume truffles with high frequency . For example,
northern flying squirrels ( Glaucomys sabrinus)’fecal
samples contains 90% over truffles spores ( Colgan,
1997; Maser et al. , 1978) . Thereare few small mam-
mals have been seen when we looked for truffles in
Yunnan forests, such as chipmunks, squirrels, flying
squirrels, rabbits and mice ( Fig. 6 ) . But there has
been no any research on this important issue in China
which is needed for better understanding and managing
the truffle forest ecosystems .
Mycorrhizas
Compared with T. melanosporum the T. indicum
complexes havemuch bigger range of host plants of
4 云 南 植 物 研 究 增刊ⅩⅥ
Fig. 3 Forests of Picea schrenkiana in Xinjiang region
where Geospora cooperi was found
Fig. 4 Natural Chinese black truffles forests
of Keteleeria evelyniana
Fig. 5 Brúle produced by Tuber indicum . Surrounding
plants are Eupatoriumodoratum
Fig. 6 Flying squirrel skin on local market,
Chuxiong, Yunnan
both broad-leaved and coniferous trees . However, only
a few ectomycorrhizal associations have been studied
and confirmed by morphological and molecular meth-
ods, such as with Pinus yunnanensis, P . armadii ,
Keteleeria evelyniana, Quercus franchetii and Castanea
mollissima (Deng, 2009) . Theectomycorrhizas formed
by the T. indicum complex are morphologically very
similar to theones produced by Tuber melanosporumin
their vertical branching emanatinghyphaeand puzzling-
like mantle structure ( Manjón et al. , 1998; García-
Montero et al. , 2008; Geng et al. , 2009; Deng,
2009) . The T. indicumcomplexwere synthesizedwith
Europenan tree species, such as Pinuspinea, Quercus
cerris, Quercus pubescens, Q. ilex and Quercus ilex
L . subsp. ballota . Themycorrhizas werevery similar to
the ones by Tuber melanosporum ( Zambonelli et al. ,
1996 , 1997; Di Massimo et al. , 1998; Manjón et al. ,
1998; García-Montero et al. , 2008) .
Kill the hen to get the eggs-truffle resources
in danger
Hundreds tonnesof Chineseblack truffles are pro-
duced annually in China now . Harvesting and trading
Chinese black truffles arebecoming amulti-milliondol-
lar industry ( Fig. 7) that has brought good income to
the local mountainous economy and farmers ( Wang et
al. , 2008 ) . Good amounts of Chinese black truffles
have been exported into European, American and Japa-
nese markets . Unfortunately, plundering commercially
harvesting driven by greed has brought big damage to
5增刊ⅩⅥ WANG andLIU: Achievements and Challenges of Rrsearch on Truffles in China
the habitats . The forest floors looked like deeply
ploughed farmsoils, tree roots werecut off and exposed
to the air after the soils weredugup for searching truf-
fles 3 - 4 times aday (Fig. 8) . The damageresulted in
sharply declining of the truffle production or no any
production at the next few years or forever ! To change
the destructive harvesting methods is quite urgent .
Careful management of truffle industry to protect truffle
resources in China is very necessary . To train truffle
dogs for hunting truffles and issue regulations for man-
agement of truffle resources has just stated in China .
Research on production of truffle seedlings and
establishment of truffleplantations Researchon produc-
ing truffle-infected seedlings for establishing plantations
has been recently carried out . The technology was ei-
ther introduced from overseas or developed by Chinese
research institutions ( Hu et al. , 2006; Lin et al. ,
2008; Geng et al. , 2009 ) . Seedlings of Pinus yun-
nanensis, P . armandii , Castanea mollissima, Quercus
franchetii , Q. aliena, Cyclobalanopsis glauca, Cory-
lus yunnanensis and Castanopsis delavayi haver been
successfully mycorrhized with T. indicumunder green-
house conditions . Q. aliena and other Chinese tree
species have also successfully been mycorrhized with
T. melanosporum . Research on technologyfor inoculat-
ing other Chinese oak species with T. melanosporum
and T. aestivum is underway .
The first truffle plantation was established on acid
soils in Taiwan in 1989 . Thesoil was limed with quick
lime (5 . 0 t?ha in 1987 and 1.7 t?ha in 1988 and 1989
respectively) to bring the soil pH up from4 .5 to 7 .0 .
Two-years-old seedlings of Cyclobalanopsis glauca in-
fected by Tuber formosanum ( = T. indicum) were
planted at a spacing of 6×6 m . Weeding was carried
out three times a year . The truffle trees were pruned
when they were four years old . No irrigation was ap-
plied . The first group of ascocarps was found in 1996
and 10 kgof truffleswereharvested from30 truffletrees
in the following year . Br?lés formed around the trees
where the ascocarps were found . Contaminated ectomy-
corrhizas have developed since 2 000 and fruiting bod-
ies of Scleroderma areolatumand Rhizopogon sp . have
been found since .
The second truffle plantation was established in
Cili County, Hunan, China, in 2002 , with T. mela-
nosporum-infected hazel ( Corylus avellana) and chest-
nut trees ( Castanea mollissima) at space of 3×3 m .
The hazel trees wereobtained fromFrance and chestnut
trees were produced fromthe Instituteof Forest Ecology
and Environment, Hunan Academy of Forestry, China .
The soil was acid and limed to raisethepH from4 .5 to
over 7 .0 . Six more truffle plantations have been estab-
lished since 2002 . One of themweas established with
Quercus aliena infected by T . indicum and T. mela-
nosporum at calcareous soils in Guiyang, Guizhou in
2004 .Two of themwere established in Panzhihua, Si-
chuan in 2007 with T. indicum infected seedlings of
Pinus armandii , Pinus yunnanensis and Castanea
mollissima, on acid soils (Fig. 9) . Threeof themwere
in Yunnan in 2008 , one of which was in Lufeng Coun-
ty , Yunnan with two years old T. indicum infected
seedlings of Quercus franchetii and Castanopsis dela-
vayi on acid red soil . The trees were produced by the
Chuxiong Forest Research Institute . Thesoil was limed
with quick lime only around the trees . Theother three
plantations have been set up in the Kunming suburb
and northwestern Guizhou andwere planted with T. in-
dicum-infected Pinus armandii and Castanea mollis-
sima (at 5×5 m) which produced fromKunming Insti-
tute of Botany, Chinese Academy of Sciences since
2007 . The soils were acid red soil and limed with
smash limestone ( around trees, 80× 80 cm) to raise
the pH to around 7 .5 . The six truffle plantations have
not produced yet except the on in Guizhou which pro-
duced its first group of ascocarps of T. indicum and
T. melanosporum in December 2008 ( Hu pers .
comm .) .
Cultivation of truffles in China is just unfolding .
However, south-west Chinahas large areasof lime-stone
soils and suitable climate for truffle growth where culti-
vation of truffles has boundless prospects (Fig. 10) .
Conclusions
(1 ) The results of researchon truffles in the last20
years revealed that China, in particular south-west China
has a huge diversity in truffle populations, genetics
6 云 南 植 物 研 究 增刊ⅩⅥ
Fig. 7 Sacs of Chinese black truffles at local truffles
market, Miyi county, Panzhihua, Sichuan
Fig. 8 Severe damaged truffle forest floor
of Pinus armandii
Fig. 9 Tuber indicumplantation established at
Panzhihua region, Sichuan in 2007
Fig. 10 Lime stone landscape,
Kunming, Yunnan
and habitats which represents akey centreof truffleor-
igin, differentiation and distribution .
(2 ) A good example of the diversity of Chinese
truffles is the extremely big variations of morphology
and genetics of T. indicum complex . Three or four
names have been proposed for this complex . DNA se-
quencings consistently divide them into two clades
(subgroups) , but these two clades can not be distin-
guished morphologically ( Wang et al. , 2006a, b;
Song, 2005; Yang et al. , 2005; Cheng, 2007;
Deng, 2009 ) . The diversity is driven by habitat and
plants diversity . In China, in particular in south-west-
ern China, the Himalayas and Hengduan Mountains
have formed since the Cenozoic Era, creating a com-
prehensive climatic conditions from tropical , subtropi-
cal , temperate to alpine cold-temperate ( including ele-
vation changing from56 to 7 700 m) and a all theveg-
etative types fromtropical rain forests to cold-temperate
coniferous forests . These areas were somehow protected
fromthe impact of the last ice age, creating an ideal
refuge for living creatures, including truffles . These
natural conditions have made this region as one of the
key centres of origin, differentiation and distributionof
tree species of Pinaceae, Fagaceae, Betulaceae, Cory-
laceae andSalicaceae ( Lu, 1999) all which areimpor-
tant host plants of truffles and therefore, the region
might be one of centres of origin, differentiation and
distributionof truffles .
( 3 ) Close biogeographic relationships between
south-western China and the Mediterranean in truffle
7增刊ⅩⅥ WANG andLIU: Achievements and Challenges of Rrsearch on Truffles in China
flora . For example, the some of truffle taxa, such as
T. aestivum, T. excavatum, T. borchii var. spherosperma
have been found in both regions . Likewise, the Tuber
indicumcomplex in south-western China, quite resem-
bles the European Périgord black truffle ( T. mela-
nosporum) (Yamanaka et al. , 2001; Chen, 2007; Je-
androz et al. , 2008) . T. pseudoexcavatumfromsouth-
western China is closely related to theEuropeanspecies
T. brumale ( Jeandroz et al. , 2008) . P . monospora is
known only from Italy and P . gigantospora only from
south-western China ( Wang et al. , 2008) .
(4) Chinese black truffles (The T. indicumcom-
plex) resemble Périgorde black truffle ( T. mela-
nosporum) not only morphologically and genetically but
also their mycorrhizas ( Zambonelli et al. , 1996 ,
1997; Di Massimo et al. , 1998; Manjón et al. ,
1998; García-Montero et al. , 2008; Geng et al. ,
2009; Deng, 2009) . They may have the same ances-
tors ( Jeandoz et al. , 2008) .
(5 ) Chineseblack truffles are important resources
for the local economy and farmers in China and also
necessary complement to deficit international truffle
market, in particular European truffle markets . Chi-
neseblack truffles are one of good edible truffles even
though their aroma is not strong as T. melanosporum
( Bellesia et al. , 2002 ) . Compared with T. mela-
nosporum, T. indicum has about the same nutritional
value and even better in thevitamin, mineral and ami-
no acid contents (Yang, 2001 ) .
(6) To change thedestructive harvesting methods
and careful management of truffle industry to protect
truffle resources in China are urgently needed .
Acknowledgement: Many thanks for thevaluable information provided
by Prof . Hu, H . T , Wang, X .H , Dr . Chen Juan and others . We appre-
ciated thatTracy Williams, Alexis Guerin and NinaWilliams corrected the
English and commented on our paper .
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9增刊ⅩⅥ WANG andLIU: Achievements and Challenges of Rrsearch on Truffles in China