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New species and distinctive geographical divergences of the genus Sparassis-fin

New species and distinctive geographical divergences of the genus Sparassis-fin
New species and distinctive geographical divergences of the genus Sparassis-fin

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Mycological Progress

ISSN 1617-416X

Mycol Progress

DOI 10.1007/s11557-012-0853-7

New species and distinctive geographical divergences of the genus Sparassis (Basidiomycota): evidence from morphological and molecular data Qi Zhao, Bang Feng, Zhu L. Yang, Yu-Cheng Dai, Zheng Wang & B. Tolgor

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ORIGINAL ARTICLE

New species and distinctive geographical divergences of the genus Sparassis (Basidiomycota):evidence from morphological and molecular data

Qi Zhao &Bang Feng &Zhu L.Yang &Yu-Cheng Dai &Zheng Wang &B.Tolgor

Received:10July 2012/Revised:8August 2012/Accepted:10August 2012#German Mycological Society and Springer 2012

Abstract Species of the genus Sparassis in East Asia were investigated using morphology and DNA sequences data.Phylogenetic analyses inferred from sequences of the inter-nal transcribed spacer (ITS),the nuclear gene coding for the ribosomal large subunit (nLSU)and partial gene coding RNA polymerase subunit II (rpb2)strongly supported line-ages corresponding to morphological features.Three taxa,S.subalpina ,S.cystidiosa f.flabelliformis and https://www.wendangku.net/doc/c511747831.html,tifolia were recognized from East Asia,and the former two taxa are new to science.The occurrence of https://www.wendangku.net/doc/c511747831.html,tifolia in Japan and in the Russian Far East was confirmed.Geographical divergences of Sparassis in the Holarctic were observed.Most species have relatively narrow distribution ranges,and taxa with intercontinental distributions were not detected.Divergence of species in the Northern Hemisphere in different clades appears to have taken place at different times:the https://www.wendangku.net/doc/c511747831.html,tifolia-crispa-radicata species complex appears to have undergone a recent radiation,while the S.subalpina-brevipes-spathularia species complex represents a relatively ancient speciation.

Keywords Cauliflower mushrooms .Morphology .Polyporales .Systematics .Taxonomy

Introduction

The Cauliflower Mushrooms (Sparassis Fr.)include species that produce flabellae with an amphigenous hymenium and a central mass giving rise to the flabellae (Desjardin et al.2004;Wang et al.2004).Members of Sparassis are distributed mainly in temperate forests,and are well known in some regions of the world (Blanco-Dios et al.2006;Burdsall and Miller 1988;Dai et al.2006;Desjardin et al.2004;Kreisel 1983).About 17names have been assigned to the genus (https://www.wendangku.net/doc/c511747831.html, ).However,due to morpho-logical plasticity and paucity,delimitation of species in the genus has been difficult.Recent molecular phylogenetic anal-yses together with morphological data recognized seven spe-cies within the genus,two of which are known only from eastern and southeastern Asia (Dai et al.2006;Desjardin et al.2004;Wang et al.2004).

During studies of macrofungi in East Asia (e.g.Du et al.2012;Feng et al.2012;Li et al.2011;Liang et al.2011;Yang 2011;Y ang et al.2012;Zeng et al.2012),rich collections of Sparassis were made.Taxonomy and biogeographic distribution of East Asia Sparassis species were revised based on morpho-logical examination and molecular phylogenetic analyses.

Materials and methods

Specimens and morphological studies Collections were obtained and photographed in the field during 1997–2011in China,the Russian Far East and Japan,and deposited in the herbarium of cryptogams of Kunming Institute of Bot-any,Chinese Academy of Sciences (HKAS),and the

Q.Zhao :B.Feng :Z.L.Yang (*)

Key Laboratory of Biodiversity and Biogeography,Kunming Institute of Botany,Chinese Academy of Sciences,Kunming 650201Yunnan,China e-mail:fungi@https://www.wendangku.net/doc/c511747831.html,

Y .-C.Dai

Institute of Applied Ecology,Chinese Academy of Sciences,Shenyang 110016Liaoning,China

Z.Wang

Department of Ecology and Evolutionary Biology,Yale University,

New Haven,CT 06511,USA

B.Tolgor

Institute of Mycology,Jilin Agriculture University,Changchun 130118Jilin,China

Mycol Progress

DOI 10.1007/s11557-012-0853-7

mycological herbarium of Jilin Agriculture University (HMJAU).Macro-morphological characters were observed from fresh specimens.For microscopical examination,dried specimens were cut and squashed in5–10%aqueous KOH. Spore sizes are given in the form of Li et al.(2011).

DNA extraction,amplification and sequencing Total DNA was extracted from silica-gel-dried or herbarium materials using the CTAB procedure of Doyle and Doyle(1987).ITS and nLSU gene fragments were amplified with primers ITS5 and ITS4(White et al.1990),and LROR and LR7(Vilgalys and Hester1990),respectively.Part of the gene encoding the second largest RNA polymerase subunit II(rpb2)was amplified with primer pairs RPB2-6F/RPB2-7R(Wang et al. 2004).PCR products were purified using the Bioteke’s Purification Kit(Bioteke Corporation,Beijing,China),and were sequenced with an ABI3730DNA analyzer and an ABI BigDye3.1terminator cycle sequencing kit(Sangon Co.,Ltd,Shanghai,China).Sequences generated in this study were submitted to GenBank;accession numbers and specimen data are provided in Table1.

Sequence alignments and phylogenetic analyses All sequen-ces were aligned with the SATéprogram(Liu et al.2009; Wang et al.2011).SATéalignments were performed with six runs of50iterations under default settings of MAFFT for alignment estimation and RAxML for phylogeny estimation using a four-core PC.The best-scored alignment was viewed with BioEdit5.0.9(Hall1999)and manually adjusted when necessary.Final concatenated alignment of three genes(ITS, nLSU and rpb2)was deposited in TreeBASE with accession no.SN11775.The phylogenetic relationships among the taxa were estimated in PAUP*version4.01(Swofford2002)under the Maximum Parsimony(MP)criterion.MP analysis was conducted using a heuristic search strategy with the following settings:the heuristic search option using the tree bisection-reconstruction algorithm.All sites were treated as unordered and equally-weighted,with gaps treated as missing data. Robustness of the internal branches of the resulting trees was tested with bootstrap analyses using1,000replications with the step-wise addition option set as simple.Grifola frondosa(Dicks.)Gray and Oligoporus rennyi(Berk.& Broome)Donk were chosen as outgroups following Wang et al.(2004).Tree scores,including tree length,consistency index[CI],retention index[RI],rescaled consistency index [RC],were also calculated.

Results

The combined ITS+nLSU+rpb2dataset consists of33 taxa and2,645aligned nucleotide sites,of which587sites (23–62,85–101,119–129,150–283,306–309,347–362,451–749,755–801,824–827,1213–1222,1336–1337, 2396–2398)were excluded due to alignment ambiguities. In the analyzed2,058nucleotides,1,427were constant, 631were variable of which164were parsimony unin-formative and467parsimony informative.Maximum parsimony analysis resulted in four equally best trees of955steps,with CI00.824,RI00.946,and RC00.779 (Fig.1).Tree topologies were,in most parts,similar to that of the most-parsimonious multi-gene tree in Wang et al.(2004).Three major clades(clades I,II,and III) were inferred.

In clade I,fourteen isolates of https://www.wendangku.net/doc/c511747831.html,tifolia Y.C.Dai& Zheng Wang from China,the Russian Far East and Japan clustered together with a98%statistic support,which shared the most recent common ancestor with S.cripsa (Wulfen)Fr.from Europe and S.radicata Weir from North America.Within the cluster of https://www.wendangku.net/doc/c511747831.html,tifolia,samples were grouped into two subclades(SW China subclade,and Northeast Asia subclade).Each subclade was statistically supported but bootstrap value for the later subclade was relatively low(62%).

In clade II,four isolates of S.subalpina formed a mono-phyletic group(bootstrap0100%)with European S.bre-vipes Krombh.and S.miniensis Blanco-Dios&Zheng Wang,and S.spathulata(Schwein.)Fr.as its sister groups (bootstrap0100%).

In clade III,three isolates of S.cystidiosa f.flabelliformis from western Yunnan,which clustered together with53% bootstrap support,formed a strongly supported clade with S. cystidiosa f.cystidiosa Desjardin et Zheng Wang(boot-strap0100%).

Taxonomy

Sparassis cystidiosa f.flabelliformis Q.Zhao,Zhu L.Yang &Y.C.Dai,f.nov.

Mycobank:MB566348

Figure2

Etymology Named for its flabelliform flabellae.

Basidioma annual,solitary,up to25cm high and30cm diam,grayish brown to yellowish brown,darkening with age,becoming brownish to light brown when dry;hyphae in the trama of the flabellum with abundant clamp connections; basidiospores(6.5)7–9(9.5)×(5.5)6–7(7.5)μm,broadly ellipsoid to subglobose,hyaline;basidia55–70×7–8.5,nar-rowly elongate-subclavate,basal septa usually with clamp connections;cystidia absent.

Holotype Q.Zhao1225(HKAS59855),2,400m,19VII. 2011,Gaoligong Mountains,Tengchong County,Yunnan Province,China.

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Basidioma(Fig.2a)annual,solitary,up to25cm high, 30cm diam,composed of a rosette of loosely arranged flabella arising from a poorly developed central core.Flabellae mostly extend from a common central mass,contorted,surface gla-brous,grayish brown to yellowish brown,darkening with age, becoming brownish to light brown when dry,top margin entire or dissected,wavy,azonate,up to10cm broad,2mm thickness.Context tough,concolorous with surface.Odor cheese-like.Taste slightly bitter.

Basidiospores(Fig.2b)[120/3/3](6.5)7–9(9.5)×(5.5) 6–7(7.5)μm(Q0(1.08)1.10–1.38(1.45),Qm01.26±0.09), broadly ellipsoid to subglobose,hyaline,thin-walled, smooth,non-amyloid,non-dextrinoid.Basidia(Fig.2c) 55–70×7–8.5μm,narrowly elongate-subclavate,4-spored,hyaline,usually with a basal clamp connection;basidioles narrowly elongate-subclavate.Subhymenium composed of tightly packed,thin-to slightly thick-walled(up to1μm thick),cylindrical or vesiculose to irregularly inflated hy-phal elements.Trama of flabellae a monomitic hyphal sys-tem composed of loosely interwoven,hyaline,inamyloid, non-gelatinous,thin-to slightly thick-walled(up to1.5μm thick),commonly clamped cylindrical hyphae(Fig.2d). Gloeoplerous hyphae present,refractive,thin-walled,flexu-ous,irregularly cylindrical to sinuous or strangulate,2–11μm diam.Cystidia absent.

Habitat and known distribution in China Terrestrial and saprophytic,solitary to scattered on the base of rotten oak

Table1Specimens used in molecular phylogenetic studies and their GenBank accession numbers

Name code Collection&herbarium no.Locality ITS nLSU rpb2

Sparassis crispa FRA ILKKA94-1587FRANCE AY218427AY218389AY218534 S.crispa GER DORISLABER GERMANY AY218442AY218404AY218544 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS55937CHINA/YN/Lijiang JN387097JN387108JN387119 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS55938CHINA/YN/Lijiang JN387098JN387109JN387120 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS55435CHINA/YN/Shangri-La JN387099JN387110JN387121 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS59358CHINA/YN/Yongping JN387100JN387111JN387122 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS59355CHINA/YN/Jianchuan JN387101JN387112JN387123 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS59356CHINA/YN/Jianchuan JN387102JN387113JN387124 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS59357CHINA/YN/Jianchuan JN387103JN387114JN387125 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS59854JAPAN/Hokkaido JQ743071JQ743081JQ743091 https://www.wendangku.net/doc/c511747831.html,tifolia HMJAU2007CHINA/JL/Jiaohe JQ743072JQ743082JQ743092 https://www.wendangku.net/doc/c511747831.html,tifolia HMJAU2955RUSSIA/Big Khekhstir JQ743073JQ743083JQ743093 https://www.wendangku.net/doc/c511747831.html,tifolia HMJAU5301CHINA/JL/Antu JQ743074JQ743084JQ743094 https://www.wendangku.net/doc/c511747831.html,tifolia(isotype)HKAS74972CHINA/JL/Antu JQ743075JQ743085JQ743095 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS74973CHINA/JL/Changchun JQ743076JQ743086JQ743096 https://www.wendangku.net/doc/c511747831.html,tifolia HKAS74974CHINA/JL/Changchun JQ743077JQ743087JQ743097 S.radicata AME TENN50232USA/TN AY218449AY218410AY218546 S.radicata CAN UBC-F12464CANADA AY218443AY218405DQ270672 S.brevipes GER ILKKA-96-1044GERMANY AY218441AY218403AY218543 S.spathulata AME7ZW-Clarku001USA/MA AY218428AY218391AY218535 S.spathulata AME8ZW-Clarku002USA/NH AY218429AY218392AY218536 S.spathulata AME11ZW-Clarku004USA/MA AY218432AY218395AY218538 S.miniensis Lou-Fungi18390SPAIN DQ270675DQ270676DQ270674 S.cystidiosa f.cystidiosa DED7410THAILAND AY256891AY256890AY256892 S.subalpina(holotype)HKAS57488CHINA/YN/Lijiang JN387093JN387104JN387115 S.subalpina HKAS57511CHINA/YN/Lijiang JN387094JN387105JN387116 S.subalpina HKAS55936CHINA/YN/Lijiang JN387095JN387106JN387117 S.subalpina HKAS55269CHINA/YN/Jianchuan JN387096JN387107JN387118 S.cystidiosa f.flabelliformis(holotype)HKAS59855CHINA/YN/Tengchong JQ743078JQ743088JQ743098 S.cystidiosa f.flabelliformis HKAS59856CHINA/YN/Tengchong JQ743079JQ743089JQ743099 S.cystidiosa f.flabelliformis HKAS59857CHINA/YN/Tengchong JQ743080JQ743090JQ743100 Grifola frondosa ZW-Clarku005USA/MA AY218415AY218413AY218521 Oligoporus rennyi KEW57Locality unknown AY218416AY287876AY218499 Mycol Progress

trees (Quercus sp.)in evergreen forest.So far only found in southwestern China (Yunnan).

Specimens examined CHINA,Yunnan Province:Tengchong County,Gaoligong Mountains,alt.2,400m,19.VII.2011,Q.Zhao 1225(HKAS 59855,holotype);the same location,19.VII.2011,Q.Zhao 1228(HKAS 59856);Tengchong

County,Houqiao,alt.2,000m,20.VII.2011,B.Feng 1070(HKAS 59857).

Notes The distinctive characters of S.cystidiosa f.flabelli-formis are the very broad flabellae with non-dissected and only slightly wavy margins,the greyish brown hymenium and relatively large basidiospores.Such traits are

very

Fig.1Phylogenetic relationships of Sparassis species inferred from a combined dataset of ITS,nLSU,and rpb2sequences.One of the four equally parsimonious trees is shown,with branch lengths proportional

to the number of mutations inferred with parsimony.Bootstrap values greater than 50%are indicated along nodes

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similar to those of S.cystidiosa f .cystidiosa .However,S.cystidiosa f .cystidiosa differs from S.cystidiosa f.flabelli-formis by the common presence of cystidia in the hymenium and its distribution in tropical southeastern Asia (Desjardin et al.2004).The sister relationship of S.cystidiosa f.fla-belliformis with S.cystidiosa f .cystidiosa was well sup-ported by the molecular phylogenetic analysis with 100%bootstrap value (Fig.1).

Sparassis subalpina Q.Zhao,Zhu L.Yang &Y .C.Dai,spec.nov.

Mycobank:MB 569412Figure 3Etymology Named for its occurrence in subalpine regions in southwestern China.

Basidioma annual,solitary,stipitate,up to 16cm high and 15cm diam,grayish when fresh,becoming brownish and leathery with age,dirty cream to light brown when dry;

hyphae in the trama of the flabellum without clamp con-nections;basidiospores (5)5.5–6.5(7)×(3.5)4–5(5.5)μm,broadly ellipsoid to subglobose,hyaline;basidia 73–85×5.5–7.5μm,narrowly elongate-subclavate,basal septa usu-ally with clamp connections.

Holotype B.Feng 759(HKAS 57488),3,050m,1.VIII.2009,Laojun Mountains,Yulong County,Yunnan Province,China.

Basidioma (Fig.3a )annual,solitary,stipitate,up to 16cm high and 15cm diam,composed of a group of flabella arising from a poorly developed central whitish stipe.Flabellae mostly extend from a common central mass,contorted,surface finely pruinose to glabrous,grayish and soft when fresh,becoming brownish and leathery with age,dirty cream to light brown when dry,1–1.5mm thick;top margin entire or slightly undulate,sometimes tooth-like or slightly laciniate,up to 2mm in thickness.Context tough,concolorous with surface.Stipe up to 7cm long,3.0

cm

Fig.2Sparassis cystidiosa f.flabelliformis (holotype).a Basidioma.b Basidiospores.c Basidia at different stage of de-velopment.d Hyphae in the trama of a flabellum

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thick at base,thinning out.Odor pleasant.Taste not recorded.

Basidiospores (Fig.3b )[120/4/4](5)5.5–6.5(7)×(3.5)4–5(5.5)μm (Q 0(1.10)1.2–1.5(1.57),Qm 01.35±0.12),broadly ellipsoid to subglobose,hyaline,thin-walled,smooth,non-amyloid,non-dextrinoid.Basidia (Fig.3c )73–85×5.5–7.5μm,narrowly elongate-subclavate,4-spored,hyaline,usually with a basal clamp connection;basidioles narrowly elongate-subclavate.Subhymenium composed of hyaline,thin-walled,tortuous,densely inter-woven hyphal segments with common presence of clamp connections.Trama of flabellum a monomitic hyphal sys-tem composed of simply septate,colorless and hyaline,thin

to thick-walled,frequently branched,interwoven hyphae 4–10μm diam;clamp connections absent (Fig.3d ).Gloeo-plerous hyphae present,refractive,thin-walled,flexuous,frequently branched,4–10μm diam.

Habitat and distribution Solitary,near the roots of living Picea likiangensis (Franch.)Pritz,Rhododendron sp.and Quercus sp.or near subalpine bamboos in northwestern Yunnan Province.So far this fungus has only been collected in subalpine Yunnan,southwestern China.

Specimens examined CHINA,Yunnan Province,Yulong County,Laojun Mountains,alt.3,050m,11.VII.2009,

B.

Fig.3Sparassis subalpina (holotype).a Basidioma.b Basidiospores.c Basidia at dif-ferent stage of development.d Hyphae in the trama of a flabellum

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Feng759(HKAS57488,holotype);the same location,11. VII.2007,Q.Zhao708(HKAS55936);the same location, 12.VII.2009,B.Feng782(HKAS57511);Jianchuan County,Jianyang,alt.3000m,29.VIII.2008,S.H.Li154 (HKAS55967).

Notes Sparassis subalpina is similar to S.brevipes,S.mini-ensis and S.spathulata by sharing the massed base of the basidioma,scattered and broad flabellae.However,the fla-bellae of S.subalpina are very broad,zonate and with only slightly contorted,thickened margins,while those of S. brevipes,originally described from Central Europe and mostly associated with Abies,Fagus and Quercus,are split into pieces that fuse with other flabellae several times below the top margin.In addition,clamp connections are common and abundant in the subhymenium but absent in the remain-ing parts of the basidioma of S.subalpina,while they are absent in all parts of S.brevipes(Wang et al.2004).Spar-assis miniensis,described based on collections made in forests of Pinus spp.from Spain,basidioma is composed of an orange to rose-purplish base,a scattered group of fragile,azonate flabellae with a strongly laciniate margin and sporadic of clamp connections in the subhymenium (Blanco-Dios et al.2006).Sparassis spathulata,originally described from eastern United States and mostly associated with Pinus spp.and Quercus spp.,produces significantly larger basidiomata,slightly longer basidiospores and no clamp connections(Burdsall and Miller1988;Desjardin et al.2004;Wang et al.2004).

Sparassis latifolia Y.C.Dai&Zheng Wang,Mycologia98: 586(2006).

Figure4

Basidioma(Fig.4a)annual,solitary,stipitate,up to 30cm high and35cm diam,composed of numerous loosely arranged flabellae.Flabellae mostly extending from a com-mon central mass,broad,dissected and slightly contorted, white to cream when fresh,becoming deep cream to shallow brown with age,pale ochraceous and light brown when dry, azonate,up to1cm broad,1mm thick,margin wavy to tooth-like.Stipe up to15cm long,2cm thick at base. Flabellar context tough,concolorous with surface.Odor pleasant.Taste indistinctly acid.

Basidiospores(Fig.4b)[340/17/17](4)4.5–5.5(6)×(3) 3.5–4(4.5)μm(Q0(1.1)1.25–1.43(1.5),Qm01.306±0.07),broadly ellipsoid to subglobose,hyaline,thin-walled,smooth.Basidia(Fig.4c)55–68×5–7μm,narrowly elongate-subclavate,4-spored,hyaline,usually with a basal clamp connection;basidioles narrowly elongate-subclavate. Subhymenium composed of delicate,hyaline,thin-walled, tortuous,densely interwoven hyphal segments with com-mon presence of clamp connections.Trama of flabellum a monomitic hyphal system composed of simply septate,col-orless and hyaline,thin to thick-walled,frequently branched,interwoven hyphae5–15μm diam;clamp con-nections common and abundant(Fig.4d).Gloeoplerous hyphae present,refractive,thin-walled,flexuous,frequently branched,7–12μm diam.

Habitat and known distribution Solitary on the base of conifers and Fagales.Distributed in eastern Russia,Japan and China.

Specimens examined CHINA,Yunnan Province,Lijiang County,Xinzhu,alt.2,900m,19.VIII.1984,Y.Xian16 (HKAS14208as“Sparassia crispa”in Zang et al.1996); Shangri-La County,Biguo,alt.3,200m,7.VII.1984,D.G. Ji76(HKAS16741as“S.crispa”in Ying and Zang1994); Yulong County,Laojun Mountains,alt.2,600m,19.VII. 2007,Q.Zhao794(HKAS55938);the same location,24. VII.2007,Q.Zhao7326(HKAS55937);Shangri-La Coun-ty,Tianchi,alt.3,200m,12.VII.2008,B.Feng324(HKAS 55435);Yongping County,Dapindi,alt.1,700m,20.VII. 2009,S.H.Li656(HKAS59358);Jianchuan County,Jia-nyang,alt.2300m,12.VII.2009,S.H.Li353,354and355 (HKAS59355,59356and59357respectively).Sichuan Province,Daocheng County,Julong,11.VIII.1984,M.S. Yuan(HKAS15728as“S.crispa”in Yuan and Sun2007). Jilin Province,Antu County,Changbaishan,14.VIII.1997, Dai2441(HKAS74972,isotype);Antu County,Edaobaihe, 20.VIII.2007,Y.Wang5301(HMJAU5301);Changchun, Jingyuetan National Forest Park,14.VII.2007,DAI12549 (HKAS74973);the same location,24.VIII.2007,DAI 10269(HKAS74974);Jiaohe County,Jiaohe,20.VIII. 2006,Y.G.Fan2007(HMJAU2007).JAPAN,Hokkaido, alt.100m,12.XI.2011,Z.L.Yang5531(HKAS59854). RUSSIA,Big khekhstir,16.VIII.2004,B.Tolgor2955 (HMJAU2955).

Notes Studies on collections from Russian Far East,Japan, northeastern and southwestern China yielded additional in-formation on https://www.wendangku.net/doc/c511747831.html,tifolia.This taxon is a common species widely distributed under trees of conifers and Fagales in the subtropic alpine and temperate regions in eastern Asia. Macroscopically,https://www.wendangku.net/doc/c511747831.html,tifolia differs from other species of Sparassis by its large,broad,dissected and slightly con-torted flabellae.Microscopically,it is characterized by the small spores and the common presence of clamp connec-tions in all tissues of the basidioma.

Sparassis latifolia and S.subalpina are similar to each other in that they both produce broad flabellae and in their ecological preference.However,https://www.wendangku.net/doc/c511747831.html,tifolia has azonate flabellae arising from a relatively more robust stipe,and the common presence of clamp connections not only in the

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subhymenium but also in other parts of the basidioma,a character shared with S.crispa and S.radicata (Dai et al.2006).

Key to the taxa of Sparassis known from East Asia and Southeast Asia

1Hyphae in the trama of the flabellum without clamp connections ............S.subalpine

1*Hyphae in the trama of the flabellum with abundant clamp connections

2Flabellae broad (up to 30cm in diam.),azonate,stipitate;hymenophore white to cream;basidiospores 4.5–5.5×3.5–4μm https://www.wendangku.net/doc/c511747831.html,tifolia

2*Flabellae broad (10–15cm in diam.),rosette;hymeno-phore dark greyish brown;basidiospores 7–9×6–7μm 3Cystidia common ............S.cystidiosa f.cystidiosa

3*Cystidia absent ............S.cystidiosa f.flabelliformis

Discussion

Biogeographic divergence in Sparassis

Supported with molecular phylogenies,10morphological taxa,S.brevipes,S.crispa ,S.cystidiosa f.cystidiosa ,S.cystidiosa f.flabelliformis ,https://www.wendangku.net/doc/c511747831.html,tifolia,S.miniensis,S.rad-icata,S.spathulata,S.subalpina ,and S.sp.(AUS31),have been recognized within Sparassis .It is interesting to note that species of Sparassis show distinct geographical distri-bution patterns,although they are primarily saprophytic (Dai et al.2006;Desjardin et al.2004;Wang et al.2004

),

Fig.4Sparassis latifolia

(HKAS55938).a Basidioma.b Basidiospores.c Basidia and basidioles at different stage of development.d Hyphae in the trama of a flabellum

Mycol Progress

living mainly on decayed wood and debris.The three taxa described in this study have only been found in East Asia so far,while the other six previously described species and one undescribed species also have a restricted distribution,i.e., S.crispa,S.brevipes and S.miniensis in Europe,and S. crispa and S.spathulata in eastern North America(One Japanese collection,CUP JA1385,regarded as S.spathu-lata,might be introduced from North America as suggested by Wang et al.2004),S.radicata in western North America, while S.cystidiosa f.cystidiosa is currently known from Thailand only,and undescribed species“S.sp.(AUS31)”only known from Tasmania,Australia(Wang et al.2004). These distribution patterns indicate that Sparassis is adapted to temperate environments with a limited number of species from tropical areas.Except for unsolved relationships be-tween European and eastern North American S.crispa,no taxa with intercontinental distributions were found to date.

The species that occur in the temperate North Hemi-sphere are clustered into two well-supported major clades (clades I and II)in the phylogeny(Fig.1),and closely related species or species pairs were found in Eurasia,East Asia-North America,or Europe-North America in each clade,indicating that broad historical exchange of Sparassis could exist in the Holarctic region.This is in accordance with observations on many ectomycorrhizal fungi,such as Boletus(Dentinger et al.2010;Feng et al.2012),Chroo-gomphus(Li et al.2009)and Amanita(Geml et al.2006, 2008;Zhang et al.2004),and many other fungi(Halling 2001;Mueller et al.2001;Petersen and Hughes2007; Redhead1989).This suggests allopatric speciation events have contributed a lot to the current distribution patterns of plants(Donoghue and Smith2004)and some fungal groups (e.g.Garnica et al.2011;Hosaka et al.2008;Vincenot et al. 2012).Furthermore,divergence of species in the temperate North Hemisphere in clades I and II appears to have oc-curred at different times based on the phylogenetic analysis (Fig.1).Further studies that include more taxa from Europe and eastern North America are critical for understanding the biogeography of Sparassis.The multilocus phylogeny sug-gest that species in Clade I(the species complex of S. latifolia-crispa-radicata)may have undergone a recent ra-diation as the three species showed limited sequence varia-tion,while the species in Clade II(the S.subalpina-brevipes-spathularia species complex)appear to represent a relatively ancient speciation event as the species in differ-ent geographical localities(East Asia,Europe,and North America)showed relatively high sequence variation. Taxonomy of https://www.wendangku.net/doc/c511747831.html,tifolia in East Asia

Sparassis latifolia was originally described from northeast-ern China(Dai et al.2006).It was originally misidentified as S.crispa in eastern Asia(Imazeki et al.1988;Mao1998,2000;Ying and Zang1994;Yuan and Sun2007;Zang et al. 1996).Dai et al.(2006)discovered that https://www.wendangku.net/doc/c511747831.html,tifolia and S. crispa can be distinguished from each other by their macro-and micro-morphological characters,i.e.,the shape of fla-bellae and the size of spores,as well as their host plant preferences.Our observations show that the long stipe of S. latifolia can also be used to separate https://www.wendangku.net/doc/c511747831.html,tifolia from S. crispa in the field.Dai et al.(2006)suggested that“S. crispa”collected and reported from far eastern Russia and Japan could be https://www.wendangku.net/doc/c511747831.html,tifolia.Studies on our collections made from the Russian Far East and northern Japan confirmed the occurrence of https://www.wendangku.net/doc/c511747831.html,tifolia in these two regions.Furthermore, analyses based on ITS sequences indicate that samples named“S.crispa”(GU138863and GU138864in GenBank) from South Korea are also closely related to https://www.wendangku.net/doc/c511747831.html,tifolia from northeast China(data not shown).These morphological and molecular data strongly indicate that https://www.wendangku.net/doc/c511747831.html,tifolia is an easily distinguishable species with a wide distribution throughout eastern Asia.

With our extensive collections from the current known distribution of https://www.wendangku.net/doc/c511747831.html,tifolia,interesting intraspecific divergen-ces of this species can be detected.As shown in Fig.1, isolates of https://www.wendangku.net/doc/c511747831.html,tifolia were divided into two subclades,the SW China subclade and the Northeast Asia subclade,with relatively high bootstrap support.Although no obvious morphological characters could be found to distinguish them,our molecular data suggests that they may represent two distinctive phylogenetic species separated by geogra-phy.Future population genetic studies should provide insights into the evolution of https://www.wendangku.net/doc/c511747831.html,tifolia. Acknowledgments This study was supported by the Natural Science Foundation of Yunnan Province(2008CD185),the Research Fund for the Large-scale Scientific Facilities of the Chinese Academy of Scien-ces(2009-LSF-GBOWS-01),and the Ministry of Science and Tech-nology of the People’s Republic of China(2008FY110300).The anonymous reviewers are gratefully acknowledged for providing con-structive comments and linguistic improvements.

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高考英语 Module 6 Old and New练习题 外研版必修3

Module 6 Old and New I. 阅读理解 A Most people who move to a foreign country or culture may go through some form of culture shock,and its degree is determined by the differences between cultures,the anxiety to adapt to a new culture and the familiarity with a new culture,etc.If you go,for example,to a culture that is far different from your own,you’re likely to experience culture shock more sharply than those who move to a new culture knowing the language and the custom of it. It is important to understand and learn how to deal with culture shock if you are to adapt successfully to your new home’s cultures.There are four general stages of cultural adjustment,and being aware of them helps you understand that culture shock won’t last long.It’s just a process you a re going through rather than a constant situation. The first stage is usually referred to as “the honeymoon stage”.Upon arriving in a new environment,you’ll be interested in the new culture.Everything will seem thrilling and everyone will seem friendly and helpful.During this stage you are merely taking in these impressions passively. But it isn’t long before the honeymoon stage gives way to the second stage—“the withdrawal stage”.The excitement you felt before is gone and problems arise.The language is hard to learn,people are unusual and unpredictable,friends,are hard to make,and simple things like shopping and going to the bank are challenges.It is at this stage that you are likely to feel anxious and homesick,and you will probably find yourself complaining about the new cu lture or country.This is the stage called “culture shock”. At some point,if you can manage it well,you’ll begin the transition into the next stage,“the recovery stage”,in which you’ll feel more confident functioning in the new culture.Customs and traditions are clearer and easier to understand.At this stage,you’ll deal with new challenges with humor rather than anxiety. 1.According to the passage,culture shock can be________. A.dealt with more easily for some people B.reduced by learning the language alone C.avoided by knowing adjustment stages D.got rid of by learning just the custom 2.“The honeymoon stage” here refers to________. A.the first month after the wedding B.the period of excitement C.the stage of adaptation D.the holiday for a newly-married couple 3.Which of the following doesn’t belong to the second stage? A.Becoming upset. B.Missing family. C.Feeling confident. D.Making complaints. 4.What would be probably discussed in the following paragraph?

高中英语Module6OldandNewSectionⅣOtherPartsoftheModule教学案外研必修3

Module 6 Old and New Section Ⅳ Other Parts of the Module [原文呈现] The Empire State Building, New York Most① of the tallest buildings in the world were built in the 1990s and in the twenty-first century, but the two tallest buildings in the US were built much②earlier. In fact, the second tallest building in the US is more than③ 75 years old! The Empire State Building, ④which was the tallest building at the time⑤ was finished in May 1931. It was the tallest building in the world until the World Trade Centre was built in New York in 1972. The World Trade Centre twin towers, which were destroyed⑥ in September 2001⑦, were 417 and 415 metres high⑧. [读文清障] ①most pron.大多数,与of连用时,句中谓语动词的单复数形式取决于of后面的名词的单复数形式。 ②much修饰比较级,用来加强语气。 ③more than多于;不仅仅是;非常 ④which was the tallest building at the time是which引导的非限制性定语从句,修饰先行词the Empire State Building。 which不能用that来替换,that不能引导非限制性定语从句。 ⑤at the time在那时 ⑥destroy v.破坏,毁坏,之所以与were连用,是因为先行词是the World Trade Centre twin towers。 ⑦which were destroyed in September 2001是which引导的非限制性定语从句。 ⑧注意此句的表达法:“...+be+数字+度量名词+adj.”表示“某物有多高/长等”。 纽约帝国大厦 [第1~2段译文] 世界上大多数最高的建筑物是在20世纪90年代和21世纪建成的,但是美国的两座最高的建筑物建成得要早得多。实际上,美国第二高的建筑物已经有超过75年的历史了! 帝国大厦是在1931年5月完工的,在那时它是最高的建筑物。它一直是世界上最高的建筑物,直到1972年世界贸易中心在纽约建成。2001年的9月被毁坏的世界贸易中心的双子塔分别有417米和415米高。 Facts about the Empire State Building:

外研版学高中ModuleOldandNew英美文化欣赏教案必修英语 解析版

【导读】《大卫·科波菲尔》是英国小说家查尔斯·狄更斯创作的长篇小说,被称为他“心中最宠爱的孩子”,于1849至1850年间,分二十个部分逐月发表。全书采用第一人称叙事,融进了作者本人的许多生活经历,反映了狄更斯希望人间充满善良正义的理想。 David Copperfieid(excerpt) That was how I was born.My early childhood was extremely happy,as my beautiful mother and kind Peggotty took care of me.But when I was about eight,a shadow passed over my happiness.My mother often went out walking,in her best clothes,with a gentleman called Mr Murdstone.He had black hair,a big black moustache and an unpleasant smile,and seemed to be very fond of my mother.But I knew that Peggotty did not like him. A few months later Peggotty told me that my mother was going to have a short holiday with some friends.Meanwhile Peggotty and I would go to stay with her brother Daniel in Yarmouth,on the east coast,for two weeks.I was very excited when we climbed into the cart,although it was sad saying goodbye to my mother.Mr Murdstone was at her shoulder,waving goodbye,as the driver called to his horse,and we drove out of the village. When we got down from the cart in Yarmouth,after our journey,Peggotty said,“That's the house,Master DaviD.” I looked all round,but could only see an old ship on the sanD.“Is that—that your brother's house?”I asked in delight.And when we reached it,I saw it had doors and windows and a chimney,just like a real house.I could not imagine a nicer place to live.Everything was clean and tidy,and smelt of fish.Now I was introduced to the Peggotty family.There was Daniel Peggotty,a kind old sailor.Although he was not married,he had adopted two orphans,who lived with him and called him Uncle.Ham Peggotty was a large young man with a gentle smile,and Emily was a beautiful,blue-eyed little girl.They all welcomed Peggotty and me warmly. 大卫·科波菲尔(节选) 就这样我来到了世上。孩提时的我非常幸福,因为有我美丽的母亲和善良的辟果提照顾我。然而,在

外研版必修三Module 6 Old and New教案设计

Module Six Old and New Step 1 Presentation Read the sentences and answer the questions. a I met a man my grandfather worked with thirty years ago. b I met a man who my grandfather worked with thirty years ago. c I wante d to visit th e house that my grandparents lived in. d Th e bus which I took back to my birth place was full o f visitors from other parts of China. 1 Do the first two sentences mean the same thing? 2 In the first two sentences, who is the subject of the verb work ----the man or the grandfather? 3 Can the words that and which be removed from the third and forth sentences without changing the meaning? Suggested answers: 1.Yes, they do 2.The grandfather 3.Yes, they can Step 2 Explanations: 定语从句的缩短 定语从句可以缩短或简化,主要通过以下形式: 一、省略作宾语的关系代词 在定语从句中作宾语的关系代词有who(m),which,that。在限制性定语从句中,当who(m)和that作宾语用于指人时,可以互换使用,通常可以省略;当that和which作宾语用于指物时,两者也可以互换使用,关系代词that/which也通常省略。例如: Is that the man(whom)/(who)/(that)you gave your tickets to?(whom,who或that可以省略) Is that the address(which)/(that)you sent the telegram to?(which或that可以省略)

四年级英语上册《Lesson 2 New and old》教案

Unit 1 The Clothes We Wear Lesson 1 Skirt and trousers 教学目标: 1.知识目标: (1)学生能听懂、会说、认读并书写词汇:is, are, new, old, it, they (2)学生能听懂、会说句子:This is a new sweater. These are old trousers. Is it new? Yes. Are they new? No. They're old. 2.能力目标: (1)在日常情境中注意观察,并用简单的英语对话(对每季度所穿衣服能加上颜色进行描述) (2)养成听录音和跟读句子的习惯。 3.情感目标: 在每天的生活中试着模仿和运用英语。 教学重点: 1.学生能听懂、会说、认读并书写词汇:is, are, new, old, it, they 2.学生能听懂、会说句子:This is a new sweater. These are old trousers. Is it new? Yes. Are they new? No. They're old. 教学难点: 代词it和they的区别及其在句子中的具体应用。 教学准备: 教学光盘、上一课的词汇卡和图片、新旧物品的图片或实物 在教室里挂一些本单元要学的服装实物,以便有真实的例子供学生练

习。 教学过程: Step1: Warm-up 1.问候Greeting。并复习学过的衣服单词,利用句型 What's this/What are these? This is a… These are… 2.课文导入Lesson hook:今天我们要学习谈论衣服的更多说法。Step 2: Presentation 1. New or old? 看第一部分4幅图,讨论左右两边的衣服有什么不同。 板书:new,old 课件出示新旧衣服图片,问:Is this a new/an old sweater? Are these new/old trousers? 出示四幅图的句子:This is a new sweater. This is an old sweater. These are new trousers. These are old trousers. 领读2遍,学生看书播放录音三遍跟读,练习读熟。 练习: (1)示范练习: 教师指向毛衣图片,与学生甲对话 What's this? It's a sweater. Is it new? Yes. It's a new sweater. 再指向裤子:What are these? These are trousers. Are these old? No. They're new trousers. (2)链式游戏:

高一英语外研版必修3教案: Module6OldandNewPeriod3

Module 6Old and New Period 3Grammar 1 & Grammar 2 & Writing 整体设计 教学内容分析 这一课时包括语法1,语法2 和写作。语法1主要是关于who 和which引导的非限制性定语从句。语法2 主要是定语从句的省略,如created by,living in。在这节课的最后一部分是发展学生的写作能力。在这一部分中,学生将要写一封关于到一个地方参观的电子邮件。 三维目标设计 Knowledge and skills 1.To help the students learn and revise the following words and expressions. Words:construction,project,region,essential,reservoir,submerge Expressions:hold back,take away,make sense,a large amount of,be full of,work with 2.To help the students learn the usage of non-defining attributive clauses and contraction of attributive clauses. 3.To help the students learn how to write an email about a visit to a place. Process and methods 1.To encourage the students to take an active part in the learning activities by giving them clear instructions on what to do while reading a new passage.In addition,correct possible mistakes they might make in a proper way by making necessary changes while repeating their sentences. 2.To encourage the students to cooperate with the others by working in pairs or in groups of four. 3.To make sure that the students pay attention to the important points by summarising the usage of non-defining attributive clauses and contraction of attributive clauses. Emotion,attitude and value 1.To let the students have confidence in learning English and express themselves freely. 2.To help the students appreciate the advantage of cooperative learning. 教学重点 1.To get the students to have knowledge of these grammar points:non-defining attributive clauses and contraction of attributive clauses. 2.To enable the students to use non-defining attributive clauses and contraction of attributive clauses. 教学难点 1.To help the students learn to use non-defining attributive clauses and contraction of attributive clauses. 2.To help the students write a description about a place. 教学过程 Lead-in 【3 min.】 Brainstorming. First ask the students to look through the following non-defining attributive clause and ask them to give their own examples. The power of the Yangtze River,which is the world's third longest river,has been harnessed by the Three Gorges Dam. Grammar 1 【8 min.】 1.Show the four sentences on page 54. a.Mao Zedong wrote a poem in which he dreamed of“walls of stone to hold back clouds and rain”. b.The Three Gorges Dam,which is the biggest construction project in China since the building of the Great Wall and the Grand Canal,has been built to control flooding. c.Sun Yat-sen,who was the leader of the 1911 Revolution,first suggested the idea in 1919. d.More than a million people who lived in the region have moved from their homes. Ask the students to read the four sentences and answer the following questions.

Module 6 Old and New 教案

Module 6 Old and New Ⅰ单元教学目标 Ⅱ目标语言

Ⅲ.教材分析与教材重组 1.教材分析 本单元的中心话题是“建筑”,所谈论的话题涉及古今中外的一些著名建筑及文化遗迹。目的在于通过单元教学,让学生了解古今中外的知名建筑,培养学生的审美观,并能就所谈论的话题写出一则新闻报道。 1.1 Introduction部分通过四个问题的提问让学生进行说前的热身,接着通过对三段小短文的展开讨论让学生明确本单元的话题。 1.2Reading部分向我们介绍了新建的三峡大坝。文章从三峡大坝兴建的原因、建成的规模、建成后的利与弊等方面进行了介绍。文章语言简洁、条理清晰,不仅有助于提高学生的阅读能力,而且在写作上也会让学生有所启发。 1.3 Grammar 1部分是学习非限制性定语从句的相关知识。

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