,2017,37(7):1404-1413 犃犮狋犪犅狅狋. 犅狅狉犲犪犾. 犗犮犮犻犱犲狀狋. 犛犻狀. :1000 4025(2017)07 1404 10 犱狅犻 :10.7606/j.issn.1000 4025.2017.07.1404 "#$%&' ()*+, 1,2, 1,2, 1,2, 3 (1 2345675689*+:;, 666303;2,< 100049;3 *=>?@ AB:;,CD 650091) -.:E ( 犘狉犻犿狌犾犪犳犾狅狉犻狀犱犪犲 ) ( 犘狉犻犿狌犾犪 )F G H I J K,L M N O H,-PE HQRSTUVFWXY (3100m) (9 4350m) Z[\]^5_`ab Lc F ;, C W 5 _,. W Z[\ ;F F,D E Z ) F, ) E FWX *F,- :(1)E L ZHQ, ZHQ, LO ZHQ, ZO\,E H J F ; ; C H ; W P W, P Z F E R Z F (2) ZF,E F,, Z, D E. F,R Z?F, HQF (3) ) E F WX * `, WXY *, O Z F! PE F*, C R )"$0BYF # $ D,%& WX #,-', 100,E FWXY WX F Z, WX () /01:MNO ;E ; Z[\; ; ) :Q948.118 23:A 犃犾狋犻狋狌犱犻狀犪犾犞犪狉犻犪狋犻狅狀狊狅犳狋犺犲犚犲狆狉狅犱狌犮狋犻狏犲犛狋狉犪狋犲犵犻犲狊狅犳犘狉犻犿狌犾犪犳犾狅狉犻狀犱犪犲 ZHANGJie 1,2,JIANG Xianfeng 1,2,CHEN Lingling 1,2,LIQingjun 3 (1KeyLaboratoryofTropicalForestEcology,XishuangbannaTropicalBotanicalGarden,ChineseAcademyofSciences,Men gla,yunnan666303,china;2universityofchineseacademyofsciences,beijing100049,china;3laboratoryofecologyand EvolutionaryBiology,StateKeyLaboratoryinConservationand UtilizationofBioresourcesin Yunnan,Yunnan University, Kunming650091,China) 犃犫狊狋狉犪犮狋 : 犘狉犻犿狌犾犪犳犾狅狉犻狀犱犪犲 isatypicaldistylousspeicesof 犘狉犻犿狌犾犪 andanendemicspeciesintheqing hai TibetPlateau.Inthisresearch,weset5plotsofnaturalpopulationfrom3100mtothetreeline(4350 m)alongthesygeramountainswithoutanthropogenicinterferenceandselectfloweringphenology,floral displaytraits,polinators,sexalocation,andnaturalseedsetascharacters.bycomparingthesecharac tersamongfivepopulationsalongaltitudinalgradients,wemakecertainlytheadaptivestrategiesofrepro :2017 03 30; :2017 06 05 :!"#$ (U1202261)!:%&(1991-),',, ()*+,- E mail:zhangjie@xtbg.ac.cn./01:,,-, ()*+,- E mail:qingjun.li@ynu.edu.cn
7 % &, :E Z[\F 1405 ductivecharacterstoaltitudinalvariation,anddiscussthepossibleinfluencesoffutureclimatechangeon distributionpaternof 犘. 犳犾狅狉犻狀犱犪犲.Wehighlightthreeconclusions:(1)initialbloomingtimeof 犘. 犳犾狅 狉犻狀犱犪犲 didnotshowacorrelationwithelevation:themiddleelevationpopulationbloomedtheearliestand thehighestpopulationbloomedlast.nevertheless,withtheincreaseofelevation,flowerlongevityand floraldisplayareaofbothmorphsof 犘. 犳犾狅狉犻狀犱犪犲 increasedsignificantly,butvarietiesofpolinatorsde creased,andboth morphstendtoalocatemoreresourcestoassurefemalesexualfunction.theadaption ofthetraitsstudiedtoheterogeneityhabitatsamongdiferentelevationsmayleadtothevariationofrepro ductivestrategies.(2)withtheincreaseofelevation,flowernumberof 犘. 犳犾狅狉犻狀犱犪犲 significantlydimin ished,butboth herbivory numberandratealsoreduced,soefectivefruitnumberofalpopulations showednodistinctdisparity,andtherewerealsonosignificantdiferencesinseednumber,showingthat byshapingcorrespondingreproductivestrategiestodiferentenvironment, 犘. 犳犾狅狉犻狀犱犪犲 guaranteedbreed ingsuccess,thenstabilizedpopulationinalelevations.(3)climatechangewilhaveprofoundandlasting impactsonthedistributionof 犘. 犳犾狅狉犻狀犱犪犲,theloweraltitudinallimitmayshiftup,butsandysoilsat higherelevationsareunfavorabletoitsgrowth,whilethetemporal spatialdistributionofpolinatorsunder climatechangearestilunknown.wepredicttheloweraltitudinallimitof 犘. 犳犾狅狉犻狀犱犪犲 wilmoveupto middleelevationandresultinadecreaseinitsdistributionoverthenext100years. 犓犲狔狑狅狉犱狊 :Qinghai TibetPlateau; 犘狉犻犿狌犾犪犳犾狅狉犻狀犱犪犲 ;altitudinalgradients;reproductivestrategies;cli matechange OV*+*+, - F*. H, / 01F* G, P* F, R L 2 [1], OV 3 4 5 6 7 8 9 : )F;. F <: =F > [2]?@,. FOV ABCPC, C F. c D ZF ) ),O Z F E [3 4] 0LF,. G* FF,G? H R,IJKLOV ZF M [5 6] N O M [7] W W [8 10] ; F O V C A *, () [11],?@, W P Q1 FRSG OV F TU [12 14] MNO VWL * LXYF G, FOV F UZ [\ 25_*. 5; G,MNO F ]H ^ 10000H, 35%G L_ P F Q [15 19] ` P M N O FSTUVVa(29 10 ~30 15 N,93 12 ~95 35 E), bzcnx, T 5? b,&, X [20] F F!& H X Y,V,L, - ;., - D M N O F R,J Wang [21] 50 NO 3 : ^ 3: F2 ; [22] R TUY,RMNO, Z2500~3500 m 30 3F 3 10 0.19, Z3500 m G 30 3F 3 10 0.25 GL ) R 100 MNO N 3 M4, 3 F O O 3 9 [23 24] F 3 MNO G* F, - G #, G Z[\Q1 \[\ ), ) F G*,- ) F WX,I J Aronne [25]. * R Z F 犘狉犻犿狌犾犪狆犪犾犻狀狌狉犻 ' R F Y, A 犘. 狆犪犾犻狀狌狉犻 F G F O 6 [26],- ( 犙狌犲狉犮狌狊狏犪狉犻犪犫犻犾犻狊 )R ) Y * WX ( 犘狉犻犿狌犾犪 ) (Primulaceae) F G, XP36, H = G F56V,G WL UZ = F WX [27] 91% FH J F J K (heterostyly, I J K distyly J Ktristyly), L J K(homostyly)Q KOI+(stigma heightdimorphism)h [28 33],I
1406 37c J KF QRG_HQ. KJ (long styled morph) K J (short styled morph) H J H JF `, KJ K KJ bp O`, KJ K KJ bp `[34 35], $ SLJ F J (disassortativemat ing) [36] L, @ ( GL P F,-,IJ. V ( 犘. 狆狅犻狊狅狀犻 )F [\F,- D: C F F [37],Brys [38] A ( 犘. 狏犲狉犻狊 )F HQ )G HQF F,HQ,HQ _., O F Z[\ F,- E ( 犘狉犻犿狌犾犪犳犾狅狉犻狀 犱犪犲 ) FIJ K,LMNO H, W X R N,R S T U V Z 3100m GWX=9 (4350 m),,- Z[\ F,- E L,-,G C W 5_,. 5_ ZO\ ; F, GY3_ :(1) Z E F * F?(2)E. F? F,( HQF?(3)MNO F ) E F W X * F? 1 R 1.1 456789:; E L. *,,.!S" #;$%&' ' Q, 1,,,()F *+,(,O 30~120cm; -. (. 15~30.,. 80.), 2 -; /, 8~10 mm,@ 0V ; 1 S, 6 S, % ' & ', 0V, 1, 2, K :13 4,_( P /, 5 6 1 3 2.5 mm *, K 13 Q 7 38; K :13 9, P / 1, 5 138 *, K 1.5mm : ( P; / S 9 [27],- E W X F N STUVL,-, E HQFW XY Z[\]^ 5_`abLc F ;( 1),B L Z 3100 m (DJMC) Z 3400 m N < (ZDNZ) Z 3750m =75> (TYFG) Z4050m 9? @ Y(LHGJT L) Z4350m 9? @ (LHGJT H) 1.2 45 1.2.1 < F?A _,-HQ B G. ( 1N ) 0L,?A CD HQF RE C H J 30, GK 0L ;,L,,-R W B3HFSFG C3.,G3. F G F? A C D 2016 6 H 2016 8H 1.2.2 =>?@A R _HQ P H JFI 30I, I P1.J F ( K,K ),G +F F B L M N (O = 0.01 mm) 5_ FF ( 2), :(1) 1 N ( 1FN,CW);(2) 1 \ ( 1 `1= F6a,CL);(3) O\ ( = 1F6a,AH);(4)K O \ ( =K F6a,SH);(5) `6 a(k = F )6a,DSA, KJ L SH AH, KJ L AH SH AL,AL L KJ F \) 1.2.3 BCDE STUV 6~8H O, C Fc R, C H R E P,?,-_ C (,-, PQ RF; _; F C (? ACD, _ ;S?A; P3d,S?A P 24h C c? A C D 2016 6H 2016 8H 1.2.4 @ RE _HQ P H JF I 30 I, I P G _ T, T B FAA U?, B. F,V a ] W X,M Y 1 ml,, Z 5 B [ \ ] 20μL M Count star B^ _,B Countstar`a^ b ^
7 % &, :E Z[\F 1407 DJMCL3100m F ;ZDNZL3400m FN <;TYFG L3750m F=75> ; LHGJT LL4050m F9? @ Y;LHGJT H L4350m F9? @ 1 E ; HQ ;WX Altitude3100mplotwassetatDJMC;Altitude3400mplotwassetatZDNZ;Altitude3750mplotwassetatTYFG; Altitude4050mplotwassetatLHGJT L;Altitude4350mplotwassetatLHGJT H Fig.1 Naturalpopulationandresearchpopulationlocationsof 犘. 犳犾狅狉犻狀犱犪犲 ZO\ KJ KJ F ( W,BSPSS22.0 U( W, B Excel2016( 2 W L KJ, L KJ CW. 1N ;CL. 1 \; SH.K O\;AH. O\;AL. \;DSA. `>\ 2 E O Long styledmorphisontheleftandshort styledmorphisonthe right.cw.corolawidth;cl.corolalength;ah.anther height;al.antherlength;sh.stigmaheight;dsa.distance betweenstigmaandanther Fig.2 Flowerdisplaytraitsmeasuredin 犘. 犳犾狅狉犻狀犱犪犲 _ TF, +^ ;AB,+^,^,. F GG 1.2.5 > R _HQ H J - 30_, -,+ ^ _ - F ( ) _ F,^ 1.3 F B Spearman F* ZO \( F W,B R W (ANOVA) 2.1 G)H < E F ZO\FF* F F 3750 m F Z H Q,6 H6,3100 m 3400 m R6H10, 3750m ; ) R3750 m, Z M, D F, 4050m ; L6H17, FL O ZFHQ,6 H 25, 19d E H JF ZO\L 0 F FF*( 犘 <0.01), Z O\F M M( 1) Z H J F R W ( 3) D ZF H J F =0, Z O\ KJ P KJ F 2.2 G)H =>?@A 1,E F. ZO\, FFF*, KJ F 1N O\ K O\ `>\ ZO\L0 FF*( 犘 <0.01), 1 \ ZO\ F( 犘 <0.05); KJ F `>\ Z O \ F F > \ ( 犘 > 0.05), ZO\ 0 FF* ( 犘 <0.01) R W ( 3) O,
1408 37c I1 "#$%&*' IJ@AH G)*K/ F Table1 Correlationanalysisbetweenelevationandcharactersofreproductivestrategiesof 犘. 犳犾狅狉犻狀犱犪犲 Item KJ Long styledmorph KJ Short styledmorph 狉犘狉犘 Flowerlongevity 0.615 0.000 0.800 0.000 1N CW 0.527 0.000 0.671 0.000 1 \ CL 0.167 0.041 0.445 0.000 AH 0.274 0.001 0.274 0.001 K O\ SH 0.327 0.000 0.405 0.000 `>\ DSA 0.217 0.008-0.142 0.084 P/O -0.204 0.012-0.262 0.000 Polendiameter 0.486 0.001 0.160 0.050 Flowernumber -0.308 0.000-0.429 0.000 Herbivorynumber -0.460 0.000-0.582 0.000 Efctivefruitnumber 0.112 0.173 0.067 0.413 Seednumber -0.160 0.050-0.031 0.709 Efctivefruitset 0.393 0.000 0.531 0.000 Herbivoryrate -0.438 0.000-0.543 0.000 : O 犘 < 0.05, O 犘 < 0.01; H JF S L150 Note: indicates 犘 < 0.05, indicates 犘 < 0.01;Totalsamplenumberofbothmorphsare150 O KJ KJ R G ZO\ ( 犘 <0.05);Y 3 ZYE KJ KJ Diferentleterswithinthesameelevationindicatesignificantdiferencebetweenlong styledandshort styled morph ( 犘 <0.05);Thesameasbelow Fig.3 Theflowerlongevityandflowerdisplaytraitsof 犘. 犳犾狅狉犻狀犱犪犲 betweenlong styledmorph andshort styledmorphineveryelevation
7 % &, :E Z[\F 1409 a. ;b. F ;c. ;d. 4 E 1 a. 犅. 狉犻犮犺犪狉犱狊犻 ;b. 犚犺犻狀犵犻犪 spp.;c. 犃狆狅狉犻犪犵狅狌狋犲犾犲犻 ;d. 犅. 犮狅狀狏犲狓狌狊 Fig.4 Polinatorsof 犘. 犳犾狅狉犻狀犱犪犲 5 ZO\E KJ KJ W Fig.5 Thesexalocationof 犘. 犳犾狅狉犻狀犱犪犲 betweenlong styledmorph andshort styledmorphineveryelevation ZF KJ KJ 1 \, Z3400m H JF 1N R @, Z H JR? @,3100 3400m F H J `>\ R, 3750m 4050m 4350m F H J `>\ 2.3 G)HBCDE E F C. Z O,3100 m Z L ( 犅狅犿犫狌狊狉犻犮犺犪狉犱狊犻 ) ( 犃狆狅狉犻犪犵狅狌狋犲犾犲犻 ) F ( 犚犺犻狀犵犻犪 spp.),3400 m Z L ( 犅狅犿犫狌狊犮狅狀狏犲狓狌狊 ) F,3750m 4050m G 4350m Z_L F, F R Z?A > R Z, F F O, ( 4), F L F C ; ZF O, F M, F F ;=
1410 37c 6 ZO\E KJ KJ Fig.6 Thenaturalseedsetof 犘. 犳犾狅狉犻狀犱犪犲 betweenlong styledmorph andshort styledmorphineveryelevation O Z4350 m, L E F P/O = C 2.5 G)H > 2.4 G)H@ 1,E KJ F 1,E H JF ZO\ 0 F F *( 犘 <0.01), (P/O) Z O \ F F ( 犘 <0.05), KJ F P/O Z O \ 0 F F F *( 犘 <0.01); KJ F ZO\ 0 F FF*( 犘 <0.01), K ZO\ F F * ( 犘 >0.05) 5 D, G ZO\ K K F, G ZO\ H J F ;, G ZO\ K F =, ZO\ H J F Z O \ F F ( 犘 > 0.05); Z O 0 FY ( 犘 <0.01), Z O 0 M( 犘 <0.01); ZO\ FF*( 犘 >0.05, 1) R W ( 6) D, Z3750m b H J, KJ KJ F O, K )P K; Z3400 m b KJ
7 % &, :E Z[\F 1411.P KJ, 4050 m b KJ.P KJ, R J D ( 6) 3 E Z O \ F, ZF3_;, O ZF _; D, D 3\ @,! *?@,E F C Fc, E bp C Fc Molau F, R ) V,? 3\, C c D, E 5, LH F ; F, H F, EH F5 [39] VWL J [40], E F P,? F, LH F Clark [41] WLF,, F Q( E F ZO \ F, ZF M,E F C., AP M, F, K J, X, K J P X, [42], K J K J F, AP, O KJ F $\( L O ) OV. AB. )F,G] 1,E F )F C, G C L,E ) Z M M,GR O Z 1L, F A F, GR AP M C F], O E LIJ K,$ S LJ F J [36], H J `>\ P, APO F W ZF*F,- D, ZO\ ME P?H, K J, P K N 1, D K J, WX Z O, KJ F KJ F, F > \ KJ O, F, D $ \ R 1 2 F O Z 6 => F,R > R, H, W F F L F $ \ [43 45] GR,E F `.P, P F ); G R, F F I [46],(G F E H JF Z O, LE ==>,O Z 1, => E Z., O, O Z, ZF,,? DE STUVa O\, WAB F* * =>?F,G HQF E AB F Z O\F* *, O \ F R, - E F W X Y 3100 m, L4350 m RMNO F Y,E FWXY *, WX! C cdf 0B E F! [47],O Z F! PE F*, C R )"$0BYF # $ D,%& WX #,?,-' E F W X = 4 ( Solomon [23] F J F, 100 MNO 3 4, Liang [48] F,- S TU V Z 100 m, 3 Y 0.66,?4 F 3 700m F Z[\ ), R 100,E FWXY WX F Z, WX F()
1412 37c LM : [1], J,,. * + [M]. <: O 3,2007:109 118. [2],.OV *+[M].<: 3,2009:1 306. [3] ARROYO M TK,ARMESTOJJ,PRIMACKRB.Commu nitystudiesinpolinationecologyinthehightemperateandes ofcentralchilei.efectoftemperatureonvisitationratesand polinationpossibilities[j]. 犘犾犪狀狋犛狔狊狋犲犿犪狋犻犮狊犪狀犱犈狏狅犾狌狋犻狅狀, 1985,149(3):187 203. [4] SARTOC M,LABOISSI?R? M C.Polinationoftwospecies of 犞犲犾狅狕犻犪 (Veloziaceae)from high altitudequartziticgrass lands,brazil[j]. 犃犮狋犪犅狅狋犪狀犻犮犪犅狉犪狊犻犾犻犮犪,2007,21(2): 325 333. [5] ARROYO M T K,ARMESTOJJ,VILLAGRAN C.Plant phenologicalpaternsinthehighandreancordileraofcentral Chile[J]. 犑狅狌狉狀犪犾狅犳犈犮狅犾狅犵狔,1981,69(1):205 223. [6] PACHECO D A,DUDLEY LS,CABEZASJ, 犲狋犪犾.Plastic responsescontributeto explaining altitudinalandtemporal variationinpotentialflowerlongevityinhighandean 犚犺狅犱狅犾犻狉 犻狅狀犿狅狀狋犪狀狌犿 [J]. 犘犔狅犛犗犖犈,2016,11(11):e0166350. [7] PENG D L,ZHANG Z Q,XU B, 犲狋犪犾.Paternsofflower morphologyandsexualsystemsinthesubnivalbeltofthe Hengduan Mountains,SW China[J]. 犃犾狆犻狀犲犅狅狋犪狀狔,2012, 122(2):65 73. [8] FABBRO T,K?RNER C.Altitudinaldiferencesinflower traitsandreproductivealocation[j]. 犉犾狅狉犪,2004,199(1): 70 81. [9] ZHAOZG,MENGJL,FANBL, 犲狋犪犾.Size dependentsex alocationin 犃犮狅狀犻狋狌犿犵狔犿狀犪狀犱狉狌犿 (Ranunculaceae):physi ologicalbasisandefectsofmaternalfamilyandenvironment. [J]. 犘犾犪狀狋犅犻狅犾狅犵狔,2008,10(6):694 703. [10],!C,,.OV F =>W [J].,2011,31(6):1157 1163. MENG L H,WANGZK,LIUCY, 犲狋犪犾.Reproductiveal locationofanalpineperennial, 犘狅犾狔犵狅狀狌犿犿犪犮狉狅狆犺狔犾狌犿 [J]. 犃犮狋犪犅狅狋犪狀犻犮犪犅狅狉犲犪犾犻 犗犮犮犻犱犲狀狋犪犾犻犪犛犻狀犻犮犪,2011,31 (6):1157 1163. [11] PAUDELBR,SHRESTHA M,BURD M, 犲狋犪犾.Coevolu tionaryelaborationofpolination relatedtraitsinanalpine ginger( 犚狅狊犮狅犲犪狆狌狉狆狌狉犲犪 )andatabanidflyinthenepalese Himalayas[J]. 犖犲狑犘犺狔狋狅犾狅犵犻狊狋,2016,211(4):1 402 1411. [12] BARRETTSC H.TheEvolution,Maintenance,andLoss ofselfincompatibilitysystems[m]//plantreproductivee cologypaternsandstrategies,new York:OxfordUniversi typress,1988. [13] BERRYPE,CALVORN.Windpolination,self Incompat ibility,andaltitudinalshiftsinpolinationsystemsinthehigh Andeangenus 犈狊狆犲犾犲狋犻犪 (Asteraceae)[J]. 犃犿犲狉犻犮犪狀犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔,1989,76(11):1602 1614. [14] ARROYO M T K,MU OZ M S,HENR?QUEZC, 犲狋犪犾. Erraticpolination,highselfinglevelsandtheircorrelatesand consequencesin an altitudinaly widespread above tree line speciesinthehigh AndesofChile[J]. 犃犮狋犪犗犲犮狅犾狅犵犻犮犪, 2006,30(2):248 257. [15],.M N O O H * F, - [J].,-,1995,17(3):233 250. WU SG,YANGYP.Onthefloraofthealpineregioninthe Qinghai Xizang (Tibet)Plateau,China[J]. 犃犮狋犪犅狅狋犪狀犻犮犪犢狌狀狀犪狀犻犮犪,1995,17(3):233 250. [16],!C."OV #$. [J].,-,2004,26(1):23 34. DENG M,ZHOU Z K.Seedplantdiversityonscreesfrom NorthwestYunnan[J]. 犃犮狋犪犅狅狋犪狀犻犮犪犢狌狀狀犪狀犻犮犪,2004,26 (1):23 34. [17] WUZY.Hengduan mountain?oraandhersignifcance[j]. 犑犪狆犪狀犲狊犲犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔,1988,63 (9):297 311. [18] %, &. V a H * F ', - [J].,-,1993,15(3):217 231. LIX W,LIJ.Apreliminaryfloristicstudyontheseedplants fromtheregionof Hengduan Mountain[J]. 犃犮狋犪犅狅狋犪狀犻犮犪犢狌狀狀犪狀犻犮犪,1993,15(3):217 231. [19] MYERSN,MITTERMEIERRA,MITTERMEIERCG, 犲狋犪犾.Biodiversityhotspotsforconservationpriorities[J]. 犖犪 狋狌狉犲,2000,403(6772):853 858. [20] LIANG E,WANG Y,XU Y, 犲狋犪犾.Growthvariationin 犃犫犻犲狊犵犲狅狉犵犲犻 var. 狊犿犻狋犺犻 alongaltitudinalgradientsinthe SygeraMountains,southeasternTibetanPlateau[J]. 犜狉犲犲狊, 2010,24(2):363 373. [21] WANGSP,WANGZH,PIAOSL, 犲狋犪犾.Regionaldifer encesinthetimingofrecentairwarmingduringthepastfour decadesin China[J]. 犛犮犻犲狀犮犲犅狌犾犲狋犻狀,2010,55 (19): 1968 1973. [22],,.M N O F ) \ [J]..,2000,45(1):98 106. YAO T D,LIU X D,WANG N L.Amplitudeofclimatic changesinqinghai TibetPlateau[J]. 犆犺犻狀犲狊犲犛犮犻犲狀犮犲犅狌犾犲 狋犻狀,2000,45(1):98 106. [23] SOLOMON S D,QIN D,MANNING M, 犲狋犪犾.Climate Change2007: The Physical Science Basis[C]//Working GroupIContributiontotheFourthAssessmentReportofthe IPCC.2007,(2):1 21. [24] KANGS,XU Y,YOU Q, 犲狋犪犾.Reviewofclimateandcry osphericchangeinthe Tibetan Plateau[J]. 犈狀狏犻狉狅狀犿犲狀狋犪犾犚犲狊犲犪狉犮犺犔犲狋犲狉狊,2010,5(1):15101. [25] ARONNE G,BUONANNO M,MICCO V D.Reproducing underawarmingclimate:longwinterfloweringandextended flowerlongevityin the only Mediterranean and maritime 犘狉犻犿狌犾犪 [J]. 犘犾犪狀狋犅犻狅犾狅犵狔,2015,17(2):535 544. [26] O 6,),,. )Y R W X ( [J].* +,2016,36(14): 4475 4484. GAO W Q,WANG X F,JIANG Z P, 犲狋犪犾.Impactof
7 % &, :E Z[\F 1413 cliamtechangeonthepotentialgeographicaldistributionpat ternanddominantcliamticfactorsof 犙狌犲狉犮狌狊狏犪狉犻犪犫犻犾犻狊 [J]. 犃犮狋犪犈犮狅犾狅犵犻犮犪犛犻狀犻犮犪,2016,36(14):4475 4484. [27] )*+,,-D..[M].<: 3,1990, 59(2):1 332. [28] WADIH AL,RICHARDSAJ.Primaryhomostylyin 犘狉犻犿 狌犾犪 L.subgenus 犛狆犺狅狀犱狔犾犻犪 (Duby)Rupr.andtheevolution ofdistylyin 犘狉犻犿狌犾犪 [J]. 犖犲狑犘犺狔狋狅犾狅犵犻狊狋,1993,124(2): 329 338. [29] RICHARDSAJ.LethalLinkageandItsRoleintheEvolution ofplantbreeding Systems[M]// Reproductive Biologyin Systematics,Conservation,and Economic Botany.Berlin: Springer,56(1):71 83. [30] VAERBAK S,ANDERSEN SB.Selectionforseedsetca pacityuncorrelated withflowerheteromorphismin 犘狉犻犿狌犾犪狏狌犾犵犪狉犻狊 Hudson[J]. 犈狌狆犺狔狋犻犮犪,1999,105(1):1 5. [31] MATSUMURA C,WASHITANII.Efectsofpopulation sizeandpolinatorlimitationonseed setof 犘狉犻犿狌犾犪狊犻犲犫狅犾犱犻 populationsinafragmentedlandscape[j]. 犈犮狅犾狅犵犻犮犪犾犚犲 狊犲犪狉犮犺,2000,15(3):307 322. [32] KATOJ,MIM.Diferencesinploidylevelsofinter specific hybrids obtained by reciprocal crosses between 犘狉犻犿狌犾犪狊犻犲犫狅犾犱犻,and 犘. 犽犻狊狅犪狀犪 [J]. 犜犺犲狅狉犲狋犻犮犪犾犪狀犱犃狆狆犾犻犲犱犌犲 狀犲狋犻犮狊,2000,101(5):690 696. [33] ENDELSP,JACQUEMYN H,BRYSR, 犲狋犪犾.Changesin pin thrumratiosinpopulationsoftheheterostyle 犘狉犻犿狌犾犪狏狌犾犵犪狉犻狊 Huds.:Doesimbalanceafectpopulation persist ence?[j]. 犉犾狅狉犪 犕狅狉狆犺狅犾狅犵狔犇犻狊狋狉犻犫狌狋犻狅狀犉狌狀犮狋犻狅狀犪犾犈犮狅犾 狅犵狔狅犳犘犾犪狀狋狊,2002,197(5):326 331. [34] DARWIN C.TheDiferentFormsofFlowersonPlantsof thesamespecies[m].john Murray,London,1877. [35] LLOYD D G,WEBB CJ.Theevolutionofheterostyly. [M ]//Evolution and Function of Heterostyly. Berlin: Springer,1992:151 178. [36] BARRETTSCH,SHOREJS.NewInsightsonHeterosty ly:comparativebiology,ecologyandgenetics[m]//self In compatibilityin FloweringPlants.Berlin:Springer,2008: 3 32. [37],/01,,.I J K [ \ F [J].*.,2015,23(6): 747 758. LIH D,RENZX,WUZK, 犲狋犪犾.Variationinfloraltraits ofdistylous 犘狉犻犿狌犾犪狆狅犻狊狊狅狀犻 (Primulaceae)alonggeograph icgradients[j]. 犅犻狅犱犻狏犲狉狊犻狋狔犛犮犻犲狀犮犲,2015,23(6):747 758. [38] BRYSR,ENDELS H, 犲狋犪犾.Therelationshipbetweenre productivesuccess and demographic structurein remnant populationsof 犘狉犻犿狌犾犪狏犲狉犻狊 [J]. 犃犮狋犪犗犲犮狅犾狅犵犻犮犪,2003,24 (6):247 253. [39] MOLAU U.Relationshipsbetweenfloweringpenologyand lifehistorystrategiesintundraplants[j]. 犃狉犮狋犻犮牔犃犾狆犻狀犲犚犲狊犲犪狉犮犺,1993,25(4):391 402. [40] D,%.6,,.O V F, - ( N.*. [J].2012,20(3):286 299. PENG D L,ZHANG Z Q,NIU Y, 犲狋犪犾.Advancesinthe studiesofreproductivestrategiesofalpineplants[j]. 犅犻狅犱犻 狏犲狉狊犻狋狔犛犮犻犲狀犮犲,2012,20(3):286 299. [41] CLARK MJ,HUSBANDBC.Plasticityandtimingofflow erclosureinresponsetopolinationin 犆犺犪犿犲狉犻狅狀犪狀犵狌狊狋犻犳狅 犾犻狌犿 (Onagraceae)[J]. 犐狀狋犲狉狀犪狋犻狅狀犪犾犑狅狌狉狀犪犾狅犳犘犾犪狀狋犛犮犻 犲狀犮犲狊,2007,168(5):619 625. [42] ZHU XF,JIANGXF,LIL, 犲狋犪犾.Asymmetricaldisassor tativepolinationinadistylousprimrose:thecomplementary rolesofbumblebeenectarrobbersandsyrphidflies[j]. 犛犮犻 犲狀狋犻犳犻犮犚犲狆狅狉狋狊,2015,5:7721 7721. [43] LLOYDDG,BAWAKS.Modificationofthegenderofseed plantsin varying conditions[j]. 犈狏狅犾狌狋犻狅狀犪狉狔犅犻狅犾狅犵狔, 1984,17(6):255 338. [44] ROFFD A.TheEvolutionofLifeHistories:TheoryandA nalysis[m].new York:Chapman & Hal,1992. [45] BRUNETJ,CHARLESWORTH D.Floralsexalocationin sequentialybloomingplants[j]. 犈狏狅犾狌狋犻狅狀,1995,49(1): 70 79. [46] RADEMAKER M CJ,DEJTJ,KLINKHAMERP G L. Polendynamicsofbumble beevisitationon 犈犮犺犻狌犿狏狌犾犵犪狉犲 [J]. 犉狌狀犮狋犻狅狀犪犾犈犮狅犾狅犵狔,1997,11(5):554 563. [47] RICHARDSJ.Primula[M].Portland:TimberPress,2014: 240 241. [48] LIANGE,BOL,ZHU L, 犲狋犪犾.Ashortnoteonlinkageof climaticrecordsbetweenarivervaleyandtheuppertimber lineinthesygera Mountains,southeastern TibetanPlateau [J]. 犌犾狅犫犪犾牔犘犾犪狀犲狋犪狉狔犆犺犪狀犵犲,2011,77(77):97 102. (:)