Kev Tshawb Fawb Txog Cistanche Hom
Mar 22, 2022
Najibeh Ataeia,b, Gerald M. Schneeweissc,⁎, Miguel Angel Garcíad,e,1, Michael Kruger, Marcus Lehnerta, Jafar Valizadehf, Dietmar Quandta,⁎
Lub koom haum Nees rau Biodiversity of Plants, Rheinische Friedrich-Wilhelms-Universität Bonn, Meckenheimer Allee 170, 53115 Bonn, Lub Tebchaws Yelemees
b General Directorate rau Agricultural Research Institute of Afghanistan (ARIA), Ministry of Agriculture, Irrigation thiab Tsiaj txhu, Badam Bagh, Kabul, Afghanistan
c Department of Botany thiab Biodiversity Research, University of Vienna, Rennweg 14, A-1030 Vienna, Austria
d Department of Biology, University of Toronto ntawm Mississauga, 3359 Mississauga, Canada
e Real Jardín Botánico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain f Department of Biology, University of Sistan thiab Baluchistan, Zahedan, Iran
Hu rau:joanna.jia@wecistanche.com/ WhatsApp: 008618081934791
Abstract
Phylogenetic kev sib raug zoo ntawm thiab nyob rau hauv non-photosynthetic parasitic kab yog notoriously paub tsis zoo, uas cuam tshuam rau peb kev nkag siab ntawm parasitic nroj tsuag. Qhov no kuj yog rooj plaub rauCistanche(Orobanchaceae), Lub Ntiaj Teb Qub genus nrog txog ob lub teb chaws, uas nws txoj kev sib raug zoo tseem tsis tau hais txog kev siv molecular phylogenetic mus kom ze. Ntawm no peb suav hais tias kev sib raug zoo ntawm phylogenetic nyob rau hauv lub genus, ntiav ib tug taxonomically thiab geographically broad qauv npog tag nrho cov yav tas los distinguished infrageneric pawg thiab feem ntau ntawm cov tam sim no paub hom. Ib qho kev sib xyaw ua ke ntawm peb cov cim plastid (trnL-trnF, suav nrog trnL intron thiab intergenic spacer (IGS), trnS-trnfM IGS, thiab psbA-trnH IGS) thiab ib lub cim nuclear (ITS) tau txheeb xyuas siv qhov siab tshaj plaws parsimony, qhov siab tshaj plaws , thiab Bayesian inference.Cistanchepoob rau hauv plaub qhov kev txhawb nqa zoo thiab thaj chaw sib txawv: East Asian Clade, Northwest African Clade, Southwest Asian Clade, thiab dav Clade. Ntawm cov no, tsuas yog East Asian Clade sib raug rau ib qho kev sau se uas tau lees paub yav dhau los, thaum lwm tus muaj cov tswv cuab ntawm ob lossis peb ntu (Widespread Clade thiab Southwest Asian Clade, ntsig txog) lossis tsis tau lees paub los ntawm taxonomic txog tam sim no (Northwest African Clade) . Whereas SouthwestAsian Clade nthuav tawm cov qauv phylogenetic muaj zog ntawm thiab ib nrab hauv cov tsiaj (East Asian Clade thiab qaum teb sab hnub poob African Clade yog monospecific), phylogenetic daws nyob rau hauv Dav Dav Clade feem ntau qis thiab cuam tshuam los ntawm qhov sib txawv ntawm cov cim nuclear thiab plastid. Ob qho tib si molecular thiab morphological cov ntaub ntawv qhia tias muaj ntau haiv neeg nyob rau hauv Cistanche yog tam sim no underestimated.
Ntsiab lus:CistancheTus tswv, Orobanchaceae, Parasite, Phylogeny, Systematics, Taxonomy
Cistanche herba Extract hmoov los ntawm Chengdu Wecistanche
1. Taw qhia
Orobanchaceae yog ib qho qauv zoo heev rau kev kawm txog kev hloov pauv ntawm cov kab mob parasitism hauv cov nroj tsuag thiab cov hauv paus phenotypic thiab kev hloov pauv caj ces (Westwood li al., 2010, Wickett et al., 2011). Lawv kuj yog ib qho piv txwv ntawm yuav ua li cas cov ntaub ntawv molecular tau ua tiav peb txoj kev nkag siab txog kev sib raug zoo ntawm phylogenetic. Raws li kev tshawb fawb txog molecular, tsev neeg kev ncig tau hloov pauv ntau thiab kev sib raug zoo ntawm kev sib raug zoo tau raug hloov kho (Wolfe li al., 2005; Bennett and Mathews, 2006; Park et al., 2008; McNeal et al., 2013; Fu et al., 2017; Li et al., 2019). Qhov dav thiab dav tshaj plaws phylogenetic tsom xam ntawm Orobanchaceae rau hnub tim yog los ntawm McNeal li al. (2013), leej twg siv cov ntaub ntawv sib xyaw ua ke ntawm tsib lub cim (nuclear: ITS, PHYA, PHYB; plastid: matK thiab rps2) suav nrog ntau dua 50 genera ntawm tsev neeg. Txawm hais tias qhov kev vam meej no, ntau yam tsis tau suav nrog hauv cov kev tshawb fawb molecular thiab nyob twj ywm unplaced (Schneeweiss, 2013). Tsis tas li ntawd, phylogenetic kev sib raug zoo nyob rau hauv genera, tshwj xeeb tshaj yog nyob rau hauv taxonomically nyuaj uas tsis yog-photosynthetic parasitic pawg, tseem nrhiav tsis tau zoo.
Ib pab pawg neeg xav tau kev tshawb nrhiav phylogenetic ntau dua yog cov tsis-photosynthetic cag-parasitic genusCistanche. Nws yog ib qho khoom plig uas muaj txiaj ntsig zoo rau kev kawm txog kev hloov pauv ntawm cov genomes loj (ntau cov chromosomes loj dua thiab sib raug zoo ntau cov genome loj dua hauv cov kab mob sib xws: Schneeweiss li al., 2004b; Weiss-Schneeweiss li al., 2006; Wicke, 20. reductive evolution ntawm plastid genomes (Li et al., 2013; Wicke et al., 2013, 2016; Liu et al., 2020), los yog hom ntau haiv neeg thiab biogeography nyob rau hauv arid cheeb tsam, tab sis ib qho ntawm cov kev tshawb fawb no yog tam sim no hampered los ntawm cov neeg pluag. kev nkag siab ntawm hom kev sib raug zoo vim tsis muaj cov ntaub ntawv phylogenetic zoo. Ob peb txoj kev tshawb fawb uas suav nrog Cistanche tsom rau kev hloov pauv ntawm plastid genome (Li et al., 2013; Wicke et al., 2013, 2016) lossis kev sib raug zoo ntawm hom siv hauv tshuaj suav tshuaj (Tomari et al., 2002, 2003 ; Han et al., 2010; Sun et al., 2012; Zheng et al., 2014), thiab yog taxonomically txwv heev. Ib yam li ntawd, cov kev tshawb fawb phylogenetic qhia nyob rau theem tsev neeg lossis tsom mus rau lwm hom, xws li Orobanche, suav nrog tsuas yog ib lossis ob peb tus neeg sawv cev ntawmCistanche(Young et al., 1999; Schneeweiss et al., 2004a; Wolfe et al., 2005; Park et al., 2008; McNeal et al., 2013; Schneeweiss, 2013; Li et al., 2019). Lawv cov txiaj ntsig qhia tau tiasCistancheyog ze ze rau lwm yam uas tsis yog-photosynthetic parasitic genera, xws li Orobanche, Phelipanche, Conopolis, los yog Epifagus (clade III ntawm McNeal li al., 2013; Orobanche-Phelipanche clade ntawm Schneeweiss, 2013), tab sis nws qhov tseeb qhov chaw tsis muaj phylo. tag nrho (Schneeweiss, 2013).
Qhov tseeb monograph ntawm tag nrho cov genus yog los ntawm Beck-Mannagetta (1930). Raws li cov duab calyx thiab tus naj npawb bracteole, nws txawv plaub ntu: (i) C. sect.Cistanchellanrog tib hom C. ridgewayana; (ii) C. sect. Ib yam khoom nrog tib homCistancheSinensis; (iii) C. sect. Heterocalyx, muaj peb hom (C. fissa, C. ambigua, thiab C. rosea); (iv) C. sect.Cistanche(nomenclatural incorrectly hu ua sect. Eucistanche los ntawm Beck Mannagetta, 1930) muaj 12 hom ntxiv. Tom qab taxonomic kev kho mob, feem ntau nyob rau hauv cov ntsiab lus ntawm lub teb chaws floras, ntxiv ob peb hom tshiab thiab ib feem hloov cov circumscriptions ntawm feem ntau morphologically variable thiab yog li taxonomically complex hom twb tau lees paub los ntawm BeckMannagetta (1930) yog li tam sim no txog 26 hom raug lees txais (Cov Ntawv Sau Npe, http://www.theplantlist.org, soj ntsuam ntawm 6. Dec. 2017). Lub genus yog dav faib nyob rau hauv cov cheeb tsam arid ntawm lub Ntiaj Teb Qub los ntawm Macaronesian Islands tuaj thiab sab hnub poob Africa mus rau nruab nrab thiab sab hnub tuaj Asia, nrog rau qhov chaw ntawm ntau hom tsiaj nyob rau sab hnub poob Asia thiab Middle East (Beck-Mannagetta, 1930). Zoo li lwm yam uas tsis yog-photosynthetic genera ntawm Orobanchaceae, Cistanche yog tus cwj pwm los ntawm kev txo qis morphological, tshwj xeeb tshaj yog cov cim vegetative (Rodrigues li al., 2011; Schneeweiss, 2013), yog li ntawd feem ntau cov cim kev kuaj mob yog los ntawm inflorescence thiab paj (zoo li paj thiab indumentum). bracts thiab bracteoles, qauv thiab indumentum ntawm lub calyx, paj xim, zoo li thiab indumentum ntawm anther), ib co ntawm cov uas tsis zoo khaws cia rau ntawm herbarium specimens. Tsis muaj kev kho taxonomic uas npog tag nrho cov hom tam sim no lees paub, cov neeg sawv cev tsis zoo hauv kev sau tshwj xeeb los ntawm cov chaw tshawb nrhiav tsawg dua, thiab qhov tsis txaus ntseeg ntawm cov cim muaj txiaj ntsig morphological ua rau muaj kev paub tsis txaus thiab tsis suav sau se ntawmCistanchehom.
Hauv txoj kev tshawb no, peb tsim kev sib raug zoo phylogenetic hauvCistancheraws li lub hauv paus rau phylogenetically kwv yees taxonomic system. Txog rau qhov kawg no, peb sau cov ntaub ntawv ua ntu zus los ntawm kev hloov pauv sai sai thiab tsim kom zoo plastid cov cim nrog rau nuclear ITS cov kab ke los ntawm kev ua piv txwv taxonomically thiab geographically comprehensive thiab txheeb xyuas cov uas siv qhov siab tshaj plaws parsimony, qhov siab tshaj plaws, thiab Bayesian txoj kev. Tshwj xeeb, peb xav kom (i) sim cov kev xav ntawm kev sib raug zoo los ntawm kev faib tawm ntawm Beck-Mannagetta (1930), piv txwv li seb nws cov tshooj yog pawg monophyletic, thiab (ii) sim seb puas yog morphologically thiab taxonomically complex hom xws li C. phelypaea thiab C. . tubulosa tsim tej pawg.
2. Cov ntaub ntawv thiab cov txheej txheem
2.1. Cov khoom cog
Ib puas-yim caum cuaj tus qauv (tsim tshiab sau, cov khoom siv herbarium lossis cov kab ke los ntawm GenBank) tau suav nrog, sib xws rau 17 yav dhau los txheeb xyuasCistanchehom ntxiv rau xya pawg neeg taxa (ib qho kev nkag ntawm Conopolis Americana, Phelipanche cf. Iberica, ob qho kev nkag ntawm Orobanche cernua, ib qho ntawm O. anatolica, O. densiflora thiab O. Transcaucasia). Ib qho piv txwv ntawmCistanchetsom rau thaj chaw dav dav rau txhua hom hauv genus. Kev txheeb xyuas hom yog raws li kev sau se siv hauv kev kho mob thiab hauv floras (Beck Mannagetta, 1930; Zhang thiab Tzvelev, 1998), tsim cov morphologically deviating hom raws li hom affinis ("aff. hom npe") lossis, qhov twg cov lus piav qhia ntawm tus tshiab subspecies yog cia siab tias (kev kho taxonomic ntawm tag nrho cov genus yog nyob rau hauv kev npaj), raws li "subsp. nov."; rau cov qauv los ntawm GenBank, uas peb tsis tuaj yeem tshawb xyuas cov ntawv pov thawj herbarium, cov ntawv sau se siv los ntawm cov thawj kws sau ntawv tau khaws cia. Cov pab pawg neeg taxa raug xaiv raws li peb cov kev paub tam sim no ntawm kev sib raug zoo hauv Orobanchaceae (Schneeweiss li al., 2004a; McNeal li al., 2013; Schneeweiss, 2013). Cov ntaub ntawv hauv zos thiab daim ntawv pov thawj suav nrog tus lej nkag mus rau GenBank tau muab rau hauv Cov Lus Ntxiv S1.
2.2. DNA cais thiab sib cais
Kev rho tawm ntawm genomic DNA los ntawm cov ntaub ntawv tshiab silica-gel qhuav corolla ua raws li CTAB-txoj cai (Doyle thiab Doyle, 1987). Feem ntau ntawm cov khoom siv herbarium raug cais los ntawm kev siv DNeasy Plant Mini Kit (Qiagen, Hilden, Lub Tebchaws Yelemees) lossis NucleoSpin Plant II (Macherey-Nagel, Düren, Lub Tebchaws Yelemees). Kwv yees li ntawm 50 mg ntawm cov khoom qhuav qhuav tau homogenized siv lub tshuab tov khoom (Retsch MM200, Haan, Lub teb chaws Yelemees) ntawm 30 Hz rau 3 min tom qab incubation hauv 700 ul extraction tsis rau yam tsawg ib teev ntawm 65 degree. Tom qab ntawd, CTABprotocol lossis cov txheej txheem muab los ntawm cov chaw tsim khoom siv tau siv.
Genomic DNA tau khaws cia ntawm −80 degree thiab dilutions tau siv rau kev nthuav dav tom qab. Peb lub cim plastid (trnL-F, suav nrog trnL intron thiab intergenic spacer (IGS), trnS-FM IGS thiab psbA-trnH IGS), nyob rau hauv thaj av loj ib qho (LSC) ntawm plastid genome, thiab nuclear Internal transcribed spacers (ITS1 thiab ITS2 ntxiv rau kev cuam tshuam 5.8S rDNA gene) raug xaiv rau phylogenetic inferences. Cov cim plastid tau rov pom dua tias ua tau zoo rau kev tshawb fawb phylogenetic ntawm qib hom (Borsch thiab Quandt, 2009). Tsis tas li ntawd, psbA-trnH twb tau siv rau DNA barcoding ntawm SuavCistanchetaxa (Han et al., 2010; Sun et al., 2012; Zheng et al., 2014). Lub cheeb tsam nuclear ITS tau raug xaiv los ua ib qho kev qhia yooj yim amenable nuclear marker tias txawm tias muaj ntau yam teeb meem (Álvarez thiab Wendel, 2003) tau ua tiav hauv ntau cov kev tshawb fawb phylogenetic nrog rau kev tsom mus rau hom kev sib raug zoo (Baldwin li al., 1995; Álvarez thiab Wendel. , 2003; Bailey li al., 2003) suav nrog Orobanche thiab lwm yam genera (xws li Schneeweiss li al., 2004a; McNeal li al., 2013). Tsis tas li ntawd, ITS2 kuj tau siv rau DNA barcoding hauvCistanche(Han et al., 2010; Sun et al., 2012; Zheng et al., 2014; Wang et al., 2018). Lub cheeb tsam trnL-F tau nthuav dav siv cov primers C thiab F (Taberlet li al. 1991). Qee qhov xwm txheej, xws li cov khoom qub thiab xav tias degraded, thaj av tau nthuav dav hauv ob qhov sib cais, piv txwv li siv primer khub C thiab D (Taberlet li al., 1991) thiab primer khub trnL460F (Worberg et al., 2007) thiab F (Taberlet et al., 1991), feem. The trnS-trnfM IGS was amplified using trnS(UAG)-pF1 (5′- ACTATACCGGTTTTCAAGGCCG-3′) and trnfM(CAU)-pR1 (5′-TAG AGC AGTTTGGTAGCTCGCA-3′; S. Wicke , Münster, pers. comm.), lub psbAtrnH IGS siv cov primers ntawm Kress li al. (2005). PCR profile rau cov cim plastid suav nrog qhov pib denaturation ntawm 5 min ntawm 94 degree, ua raws li 30 cycles txhua nrog 1 min ntawm 94 degree, 1 min ntawm 55 degree, 90 s ntawm 72 degree, thiab qhov kawg elongation kauj ruam ntawm 7 min ntawm 72 degree. Lub cheeb tsam ITS tau nthuav dav siv cov primers ITS4 thiab ITS5 (Dawb li al., 1990) nrog rau kev nthuav dav ntawm 5 min ntawm 94 degree tom qab los ntawm 40 cycles txhua nrog 1 min ntawm 94 degree, 1 min ntawm 48 degree nrog lub sijhawm- nce ntxiv ntawm 4 s ib lub voj voog, 45 s ntawm 68 degree, thiab cov kauj ruam kawg txuas ntxiv ntawm 7 min ntawm 68 degree. Cov PCRs tsis ua tiav tau rov qab siv cov primers sab hauv 5.8S106-R (5′- AGGCGCA ACTTGCGTTCAAA -3′) thiab 5.8S32-F (5′-GCATCGATGAAGAGAGAT AGC-3′; D. l. Nickrent, Southern Illinois University, USA, pers. comm.) ua ke nrog cov primers sab nraud ITS5 thiab ITS4. PCR cov tshuaj tiv thaiv tau ua nyob rau hauv ib ntim ntawm 25 ul thiab suav nrog 1.5 UGoTaq Flexi DNA polymerase (Promega, Madison, USA), 0–0.2 M betaine monohydrate, 0.4 µM ntawm txhua tus pem hauv ntej thiab rov qab primer, 0.15 mM dNTPs (Carl Roth, Karlsruhe, Lub teb chaws Yelemees), 1 mM MgCl2 nyob rau hauv 1xGoTaq Flexi tsis, 1 ul genomic DNA ntawm tsis paub concentration, thiab dej. Rau qee cov khoom siv herbarium degraded heev Npaj-To-Go PCRBeads (Amersham-Pharmacia Biotech, Amersham, UK) tau siv ua raws li cov chaw tsim khoom cov lus qhia. Raws li feem ntau xav tau rau DNA-kev cais ntawm cov khoom siv herbarium, PCR additives xws li 1 ul PVP-40(10–40 feem pua) thiab/los yog 5 ul enhancer tov P (5 ×; PeqLab, Erlangen, Germany) tau ntxiv rau cov tshuaj tiv thaiv ntawm tus nqi dej. Amplification cov khoom tau gel-purified ntawm 1 feem pua agarose gel siv cov khoom siv thePeqLab PCR purification kit (Peqlab) lossis cov khoom siv ceev ceev PCR purification (Qiagen). Rau nested ITS-PCR cov khoom ntawm <300 bp="" (kev="" ua="" kom="" zoo="" dua="" los="" ntawm="" cov="" khoom="" siv="" herbarium,="" uas="" feem="" ntau="" muaj="" cov="" kab="" mob="" fungi),="" gel="" concentration="" ntau="" dua="" (1.4="" feem="" pua)="" thiab="" lub="" sijhawm="" ua="" haujlwm="" ntev="" dua="" raug="" xaiv.="" cov="" khoom="" siv="" pcr="" huv="" si="" tau="" ua="" tiav="" los="" ntawm="" macrogen="" (seoul,="" kaus="" lim="" kauslim)="" nrog="" rau="" kev="" ua="" kom="" nrov="" nrov="" thiab="" cov="" primers="" sab="" hauv="" ntxiv="" tau="" hais="" los="" saum="" no,="" qhov="" xav="">
cistanche wrkung
2.3. Kev sib xyaw ua ke, indel coding, thiab phylogenetic tsom xam
DNA sequences tau sib sau ua ke thiab ua raws li PhyDE 0.97 (Muller li al., 2006). Tom qab Olsson et al. (2009), thaj tsam ntawm homology tsis paub meej (mutational hotspots) tau sau tseg hauv PhyDE thiab tshem tawm ntawm kev tshuaj ntsuam. Cov kev hloov pauv tau raug cais tawm hauv qhov kev sib tw thiab suav nrog kev rov qab-ua tiav raws li tau hais dhau los (Quandt li al., 2003; Borsch thiab Quandt; 2009). Indels tau coded siv yooj yim indel coding (SIC; Simmons thiab Ochoterena, 2000) raws li siv nyob rau hauv SeqState 1.25 (Müller, 2005) thiab ntxiv raws li ib tug ntxiv cov ntaub ntawv muab faib. Nyob rau hauv tag nrho cov ntaub ntawv matrices, ib ntus qhov sib txawv tau raug kho raws li cov ntaub ntawv uas ploj lawm thiab cov haujlwm sib luag tau raug kho raws li qhov hnyav sib npaug.
Kev tshuaj xyuas tau ua los ntawm peb cov ntaub ntawv sib txuas ua ke sib txuas nrog cov cim plastid concatenated, ITS, thiab cov ntaub ntawv sib xyaw ua ke (plastid thiab nuclear markers ua ke), txhua tus nrog lossis tsis muaj indels, uas ua rau tag nrho ntawm 6 cov ntaub ntawv tau txheeb xyuas. Ua ntej qhov kev tshuaj ntsuam ua ke, ib qho cim cov ntaub ntawv teev tau raug tshuaj xyuas rau qhov tsis sib haum xeeb ntawm cov cim los ntawm kev txheeb xyuas cais ntawm txhua qhov chaw hauv MrBayes siv qhov chaw pib (cov ntaub ntawv tsis pom). Qhov siab tshaj plaws parsimony tsom xam (MP) ntawm concatenated nucleotide matrix nrog thiab tsis muaj qhov txuas ntxiv indel matrix tau ua nyob rau hauv PAUP* 4.0b10 (Swofford, 1999) siv parsimony ratchet (Nixon, 1999) ntawm cov ntaub ntawv hais kom ua. los ntawm PRAP2 (Muller, 2004). Cov kev teeb tsa ratchet nram qab no tau siv: 200 iterations nrog 25 feem pua ntawm cov haujlwm randomly up-weighted (hnyav=2) nyob rau hauv txhua replicate thiab 10 random ntxiv cycles. Qhov siab tshaj plaws (ML) tsom xam ntawm concatenated nucleotide matrix
nrog thiab tsis muaj qhov txuas ntxiv indel matrix, tau ua los ntawm RAxML 8 (Stamatakis, 2014) siv tus qauv GTRCAT. Rau ob qho MP thiab ML kev txheeb xyuas kev txhawb nqa tau kwv yees los ntawm bootstrapping (Felsenstein, 1985) siv 10, 000 replicates. Bayesian inferences (BI), ua rau tag nrho rau cov ntaub ntawv teev tseg (ib leeg thiab cov cim sib txuas, nrog thiab tsis muaj indels coded) tau ua nrog MrBayes 3.2.2 (Huelsenbeck thiab Ronquist, 2001; Ronquist thiab Huelsenbeck, 2003) Γ ntxiv rau GTR ntxiv. Kuv ua qauv rau cov nucleotide muab faib (ob qho qauv tsis sib txuas hauv cov ntaub ntawv sib xyaw ua ke) thiab cov qauv txwv qhov chaw (F81-zoo li; Felsenstein, 1981) rau qhov muab faib indel nrog rau qhov ua ntej. Rau kev khiav nrog 10 lab tiam txhua thiab plaub chains txhua tus (ib qho txias thiab peb lub tshuab cua sov uas siv cov cua sov ua ntej) tau khiav hauv qhov sib npaug, piv txwv txhua 1000 tiam. Thawj 25 feem pua tau muab pov tseg raws li kev kub hnyiab, uas yog qhov zoo tom qab chains tau mus txog qhov chaw nyob, raws li tau teev tseg hauv Tracer v.1.7.1 (Rambaut li al., 2018). Cov ntoo raug kho siv TreeGraph2 (Stöver thiab Müller, 2010).
3. Cov txiaj ntsig
Cov yam ntxwv ntawm cov ntaub ntawv tag nrho, qhov twg tsis muaj txoj haujlwm raug tshem tawm, thiab ntawm cov ntaub ntawv teeb tsa, qhov twg hotspots (rau cov ntsiab lus ntawm cov saib Cov Lus Ntxiv S2) tau raug tshem tawm (mononucleotide rov ua dua) lossis rov qab (hais plaub hau cuam tshuam inversions), tau muab rau hauv Table 1. Feem ntau cov indels hauv cov ntaub ntawv tau rov ua dua ntawm cov kab uas nyob ib sab (piv txwv li, cov kab ke yooj yim rov ua dua [SSR]), qhov ntev ntawm 2 txog 20 nucleotides (cov ntaub ntawv tsis qhia). Tsis tas li ntawd, hauv psbA-trnH spacer, kev tshem tawm ntawm 1148 bp nyob rau hauv cov matrix sib raug zoo tau pom hauv C. aff. fissa 2, coj mus rau qhov txo qis hauv ntu ntev mus rau 20 feem pua ntawm qhov nruab nrab ntev.
Peb pom tsis muaj qhov tsis sib haum xeeb tseem ceeb ntawm cov kev sib raug zoo ntawm phylogenetic inferred los ntawm ib qho plastid cov cim (cov ntaub ntawv tsis qhia), tab sis ntawm concatenated plastid markers thiab nuclear marker (Supplementary Fig. S1–S2). Cov no feem ntau pom muaj nyob rau hauv clade nrog rau, ntawm lwm tus, C. phelypaea thiabCistanchetubulosa (piv txwv li, Txoj Cai Dav Dav: saib kab lus tom ntej), uas yog tus cwj pwm los ntawm qhov tob tob ntawm qhov sib txawv (saib phylogram hauv daim duab 1). Tsis muaj qhov sib txawv tseem ceeb ntawm cov ntoo uas tau lees paub los ntawm cov ntaub ntawv teev nrog lossis tsis muaj coding indels; Txawm li cas los xij, kev siv cov ntaub ntawv indel tau nce tag nrho cov kev daws teeb meem topological, qhov uas nws cuam tshuam rau cov txiaj ntsig ntawm kev txhawb nqa yog qhov tsis meej (Daim duab 1, Cov duab ntxiv. S1–S3).
Cov ntoo phylogenetic tau los ntawm ML thiab BI raws li kev sib xyaw plastid thiab cov cim nuclear, ob qho tib si nrog indels (ML log-zoo li tus qhab nia ntawm −21272.719; harmonic txhais tau tias ntawm log-zoo li cov qhab nia los ntawm BI ntawm −21432.751) thiab tsis muaj indels (ML log-zoo li tau qhab nia ntawm −17135.239; harmonic txhais tau tias ntawm log-zoo li cov qhab nia los ntawm BI ntawm −17654.075), tau congruent thiab yielded ib tsob ntoo zoo daws teeb meem (Fig. 1). Hauv qab no, peb yuav tsom mus rau cov txiaj ntsig los ntawm kev sib xyaw ua ke, xa mus rau ib qho kev txheeb xyuas ib leeg (concatenated plastid piv rau nuclear) tsuas yog thaum muaj qhov sib txawv tseem ceeb. Cov kev txheeb xyuas ua ke tau txheeb xyuasCistancheraws li ib pawg monophyletic, txawm hais tias tsuas yog tsis muaj zog txhawb nqa (ML 61/ < 50;="" bi="" 0.77/0.91;="" txhawb="" cov="" txiaj="" ntsig="" los="" ntawm="" kev="" txheeb="" xyuas="" nrog="" coded="" indels="" tsis="" muaj="" coded="" indels).="" hauv="" mp="" tsom,="" txoj="" hauj="" lwm="" ntawm="" c.="" sinensis="" nrog="" rau="" lwm="">Cistanchehom thiab cov se tawm ntawm pawg neeg tsis tau daws (Ntxiv Fig. S3).Cistanchepoob rau hauv plaub qhov loj-txhawb clades (Fig. 1, Supplementary Fig. S3), tam sim no hu ua East Asian Clade, Southwest Asian Clade, Northwest African Clade, thiab Dav Clade, feem. Sab hnub tuaj-Asian Clade, tsuas muaj C. Sinensis (MP 100/100; ML 100/100; BI1.00/1.00) los ntawm Tuam Tshoj thiab Mongolia, raug suav hais tias yog tus muam rau cov seem ntawmCistanche(MP 100/100; ML 100/100; BI 1.00/1.{23}}). Sab hnub poob Asian Clade (MP 100/100; ML 100/100; BI 1.00/1.00), muaj ntau hom tsiaj los ntawm sab hnub poob mus rau Central Asia, yog viv ncaus rau lub clade (MP 100/100; ML 100/100; BI 1.00/1.00) suav nrog rau sab qaum teb sab hnub poob African Clade thiab Widespread Clade. Whereas lub Northwest African Clade (MP 100/100; ML 100/100; BI 1.00/1.00) tsuas muaj ib hom txwv rau qaum teb hnub poob teb chaws Africa, qhov dav Clade (MP 100/100; ML 100/100; BI 1.00 / 1.00). txog kaum hom sib koom ua ke muaj kev faib tawm thoob plaws hauv Atlantic ntug dej hiav txwv ntawm Europe thiab Africa mus rau Central Asia.
Hauv Southwest Asian Clade, cov qauv phylogenetic tau hais tawm (Daim duab 1, Ntxiv Fig. S3). Clade A (MP 98/100; ML 100/ 100; BI 1.00/1.{23}}0) suav nrog tsuas yog C. ambigua, uas poob rau hauv ob thaj chaw sib cais subclades, ib qho (MP 79/1{{30}}}; ML 75/ < 50;="" bi="" 1.00="">< 50)="" uas="" muaj="" kev="" nkag="" mus="" rau="" sab="" qaum="" teb="" sab="" hnub="" tuaj="" iran,="" qhov="" thib="" ob="" (mp="" 75/51;="" ml="" 82/="">< 50;="" bi="" 1.00/0.="" 51)="" nrog="" qee="" qhov="" kev="" nkag="" los="" ntawm="" sab="" qaum="" teb="" iran="" nrog="" rau="" ob="" txoj="" kev="" nkag="" los="" ntawm="" sab="" hnub="" poob="" afghanistan.="" clade="" a="" yog="" tus="" muam="" (mp="" 70/52;="" ml="" 69/="">< 50;="" bi="" 1.00/0.93)="" rau="" c.="" ridgewayana="" los="" ntawm="" afghanistan.="" cov="" viv="" ncaus="" tom="" ntej="" (mp="" 98/99;="" ml="" 100/100;="" bi="" 1.00/1.00)="" yog="" clade="" b="" (mp="" 62/68="" ;="" ml="" 75/79;="" bi="" 0.97/1.00)="" comprises="" the="" accessions="" of="" c.="" aff.="" ridgewayana.="" nyob="" rau="" hauv="" clade="" b="" phylogenetic="" qauv="" sib="" haum="" mus="" rau="" geography,="" vim="" hais="" tias="" accessions="" los="" ntawm="" qaum="" teb="" hnub="" poob="" thiab="" central="" iran="" (aff.="" ridgewayana="" 2)="" tsim="" ib="" tug="" subclade="" nested="" nyob="" rau="" hauv="" ib="" qeb="" ntawm="" accessions="" los="" ntawm="" feem="" ntau="" central="" thiab="" yav="" qab="" teb="" iran="" (aff.="" ridgewayana="" 1).="" kev="" sib="" raug="" zoo="" ntawm="" cov="" kev="" nkag="" ntawm="" c.="" ridgewayana="" sl="" (c.="" ridgewayana="" thiab="" c.="" aff.="" ridgewayana)="" txawv,="" txawm="" li="" cas="" los="" xij,="" ntawm="" cov="" cim:="" whereas="" c.="" ridgewayana="" sl="" tau="" inferred="" li="" monophyletic="" yam="" tsis="" muaj="" kev="" txhawb="" nqa="" sab="" hauv="" los="" ntawm="" its="" cov="" ntaub="" ntawv="" (supplementary="" fig.="" s2)="" ,="" nws="" tau="" inferred="" li="" paraphyletic="" los="" ntawm="" plastid="" cov="" ntaub="" ntawv="" nrog,="" piv="" rau="" cov="" kev="" tsom="" xam="" ua="" ke,="" hloov="" txoj="" hauj="" lwm="" ntawm="" c.="" aff.="" ridgewayana="" 1="" thiab="" c.="" aff.="" ridgewayana="" 2.="" cov="" viv="" ncaus="" tom="" ntej="" mus="" rau="" clade="" suav="" nrog="" c.="" ambigua="" thiab="" c.="" ridgewayana="" sl="" (mp="" 74/60;="" ml="" 96/93;="" bi="" 1.00/1.{102}})="" yog="" clade="" c="" (mp="" 1{104}/100;="" ml="" 100/100;="" bi="" 1.00/1.00)="" uas="" muaj="" c.="" aff.="" fissa="" 1="" los="" ntawm="" afghanistan.="" cov="" viv="" ncaus="" tom="" ntej="" no="" yog="" (i)="" ib="" leeg="" nkag="" ntawm="" c.="" aff.="" fissa="" 2="" los="" ntawm="" azerbaijan="" (mp="" 57/100;="" ml="" 84/85;="" bi="" 0.88/0.57),="" (ii)="" clade="" d="" (mp="" 100/100;="" ml="" 100/100;="" bi="" 1.00/1.00)="" suav="" nrog="" c.="" salsa="" los="" ntawm="" sab="" hnub="" poob="" asia="" thiab="" los="" ntawm="" tuam="" tshoj="" (mp="" 100/100;="" ml="" 80/90;="" bi="" 0.99/0.99),="" thiab="" (iii)="" clade="" e="" (mp="" 100/100;="" ml="" 100/100;="" bi="" 1.00="" 1.00)="">Cistanchedeserticola los ntawm Tuam Tshoj thiab Mongolia (MP 100/100; ML 100/100; BI 1.00/1.00).
Fig. 2. Tus yam ntxwv paj thiab calyx cov duab rau cov tswv cuab ntawm East Asian Clade (J), Southwest Asian Clade (F–I), Northwest African Clade (K), thiab dav Clade (A–E, L– M) ntawm Cistanche. (A) C. senegalensis, (B) C. flava, (C) C. tubulosa subsp. tubulosa, (D) C. violacea, (E) C. phelypaea subsp. nov., (F) C. fissa, (G) C. salsa, (H) C. ridgewayana, (I) C. ambigua, (J) C. Sinensis, (K) C. spec. nov., (L) C. rosea, (M) C. laxiflora.
Nyob rau hauv dav Clade kev sib raug zoo muaj qee zaus tsis meej vim tias tsis muaj kev daws teeb meem ntawm cov ntaub ntawv nuclear nrog rau qhov tsis sib xws ntawm cov cim plastid thiab nuclear (Supplementary Fig. S1–S2). Cov kev txhawj xeeb no tsis yog txoj haujlwm ntawm ib leeg nkag mus (xws li, C. Senegalese ED1096,Cistanchetubulosa sub. tubulosa ED891) tab sis kuj tseem muaj cov kab mob ntawm cov subclades loj dua. Piv txwv li, whereas plastid cov ntaub ntawv inferred C. Rosea raws li ib tug zoo-paub viv ncaus rau lwm yam tsiaj (BI 1.00; Supplementary Fig. S1), cov ntaub ntawv nuclear qhia tias nws yog ib tug tswv cuab ntawm ib tug clade (BI 1. 00) suav nrog txhua hom tshwj tsis yog C. phelypaea, C. violacea, thiab feem ntau nkag mus ntawm C. lutea (Supplementary Fig. S2). Yog li ntawd, kev sib raug zoo ntawm cov clades loj (clades F–J) inferred los ntawm cov kev ntsuam xyuas ua ke feem ntau daws tsis zoo thiab tsis zoo txhawb nqa (Daim duab 1). Qhov kev zam tsuas yog clade F (MP 100/100; ML 100/100; BI 1.00/1.00), muaj C. rosea los ntawm Arabian Peninsula (hauv kev tshuaj xyuas ITS, qhov sib thooj thiab tsuas yog tsis muaj zog txaus txhawb nqa clade ntxiv nrog rau kev nkag mus rau ED1096 ntawm C. senegalensis: Ntxiv Fig. S2), uas nws tus muam-pab kev sib raug zoo rau clade (MP 99/94; ML 98/99; PP 1.00/ 0.99) suav nrog cov tsiaj seem (clades G–J; Fig. 1) tau txais kev txhawb nqa zoo.
herba cistanches
Clade G (MP 63/99; ML 84/54; BI 1.00/ < 50)="" muaj="" c.="" laxiflorafrom="" iran="" thiab="" afghanistan="" ua="" ke="" nrog="" ob="" qhov="" kev="" nkag="" ntawm="" c.="" tubules="" los="" ntawm="" tuam="" tshoj="" (gb1="" thiab="" gb2).="" nyob="" rau="" hauv="" clade="" g,="" c.="" laxiflora="" yog="" paraphyletic,="" vim="" hais="" tias="" lub="" subclade="" ntawm="" ob="" c.="" tubulosa="" accessions="" (mp="" 98/65;="" ml="" 99/73;="" bi="" 1.00/0.77)="" grouped="" hauv="" a="" clade="" (mp="" 54/="">< 50;="" ml="" 60/91;="" bi="" 1.00/1.{24}})="" nrog="" rau="" kev="" nkag="" ntawm="" c.="" laxiflora="" los="" ntawm="" sab="" qaum="" teb="" thiab="" sab="" qab="" teb="" iran="" nrog="" rau="" afghanistan="" (mp="" 85="">< 50;="" ml="">< 50/50;="" bi="">< 0.50="" 0.74)="" rau="" kev="" cais="" tawm="" ntawm="" kev="" nkag="" ntawm="" c.="" laxiflora="" los="" ntawm="" central="" iran="" tsim="" ib="" qho="" clade="" cais="" (mp="" 99/95;="" ml="" 99/95;="" bi="" 1.="">
Clade H (MP unsupported; ML 97/50; PP 1.00/ < 50)="" muaj="" c.="" senegalensis="" (sab="" hnub="" tuaj="" teb="" chaws="" africa)="" thiab,="" nested="" nyob="" rau="" hauv,="" ib="" tug="" monophyletic,="" tseem="" unsupported,="" c.="" flava="" (southwestern="" asia)="" thiab="" subclade="" (mp="" 90/73;="" ml="" 86/69;="" bi="" 0.79/0.{12}})="" muaj="" xws="" li="" accessions="" ntawm="" c.="" aff.="" tubulosa="" los="" ntawm="" sab="" qab="" teb="" arabian="" peninsula="" (oman,="" yemen).="" cov="" ntaub="" ntawv="" pov="" thawj="" rau="" clade="" h="" tuaj="" qhov="" tseem="" ceeb="" los="" ntawm="" cov="" ntaub="" ntawv="" plastid="" ib="" leeg="" vim="" tias="" hauv="" its="" c.="" flava="" tau="" suav="" tias="" yog="" ze="" ze="" rau="" c.="" laxiflora="" thiab="" kev="" sib="" raug="" zoo="" ntawm="" c.="" senegalensis="" thiab="" c.="" tubulosa="" tseem="" tsis="" tau="" daws="" (ntxiv="" fig.="" s2).="" hauv="" clade="" h,="" tsis="" tsuas="" yog="" kev="" sib="" raug="" zoo="" ntawm="" c.="" tubulosa="" thiab="" c.="" flava="" rau="" c.="" senegalensis="" tau="" daws="" tsis="" zoo="" thiab="" kev="" txhawb="" nqa="" tsis="" zoo,="" tab="" sis="" kuj="" yog="" cov="" kab="" hauv="" c.="" senegalensis="" thiab="" hauv="" c.="" flava,="" tsuas="" yog="" qhov="" tshwj="" xeeb="" yog="" subclade="" ntawm="" plaub="" c.="" .="" flava="" accessions="" (flava="" subsp.="" nov.)="" los="" ntawm="" qaum="" teb="" iran="" (mp="" 100/97;="" ml="" 100/99;="" bi="">
Clade I (MP unsupported; ML 62/54; PP 1.00/ < 0.50)="" muaj="" cov="" khoom="" tshwj="" xeeb="" ntawm="" c.="" tubulosa,="" npog="" thaj="" chaw="" dav="" dav="" ntawm="" cape="" verde="" islands="" thiab="" mali="" ntawm="" lub="" arabian="" peninsula="" thiab="" iran="" mus="" rau="" pakistan="" thiab="" is="" nrias="" teb.="" cov="" ntaub="" ntawv="" pov="" thawj="" rau="" clade="" h="" kuj="" tseem="" tuaj="" yeem="" los="" ntawm="" cov="" ntaub="" ntawv="" plastid="" ib="" leeg,="" vim="" tias="" hauv="" its="" pawg="" no="" tau="" suav="" tias="" yog="" paraphyletic="" thiab="" feem="" ntau="" tsis="" tau="" daws="" nrog="" c.="" rosea="" (clade="" f),="" c.laxiflora="" (clade="" g)="" thiab="" c.="" senegalensis="" thiab="" c.="" .="" flava="" (clade="" h).="" phylogenetic="" qauv="" hauv="" clade="" kuv="" feem="" ntau="" tsis="" muaj="" zog="" thiab="" tsis="" pom="" qhov="" txawv="" ntawm="" thaj="">
Clade J (MP 92/53; ML 97/87; PP 1.00/0.99) muaj C. lutea sl (ie, suav nrog C. lutea, C. aff. lutea 1 thiab C. aff. lutea 2; Macaronesian Islands thiab Mediterranean) thiab, zes nyob rau hauv, C. Brunner (western Africa), C. phelypaea (Macaronesian Islands, Atlantic thiab Mediterranean ntug dej hiav txwv ntawm Africa thiab sab qab teb Europe), thiab C. violacea ( North Africa).Cistanchephelypaea (MP 100/98; ML 100/ 100; BI 1.00/1.{{ 31}}0) tsim, ua ke nrog qib ntawm C. lutea sl (accessions los ntawm tag nrho cov nyob rau hauv no hom kev faib ntau yam) ntxiv rau ob accessions ntawm C. violaceus (ED807 thiab ED1{{ 41}}12 los ntawm Saudi Arabia thiab Jordan, feem) thiab hybrid ntawm C. aff. lutea 1 thiab C. violacea, ib tug tsis muaj zog txhawb clade (MP 55/-; ML 76/67; BI 0.64/ < 0.50)="" uas="" yog="" ib="" tug="" muam="" (mp="" 90/53;="" ml="" 97/92;="" bi="" 0.96/0.89)="" rau="" ib="" lub="" clade="" (mp="" 68/58;="" ml="" 93/82;="" bi="" 0.90/0.63)="" suav="" nrog="" tag="" nrho="" tab="" sis="" ob="" qho="" dhau="" los="" hais="" txog="" kev="" nkag="" ntawm="" c.="" violacea.="" nyob="" rau="" hauv="" sib="" piv="" rau="" c.="" violacea,="" qhov="" twg="" thaj="" chaw="" sib="" cais="" mus="" rau="" hauv="" ib="" tug="" dav="" dav="" clade="" (mp="" 53/70;="" ml="" 84/85;="" bi="" 0.92="" 0.59)="" thiab="" ib="" qho="" tshwj="" xeeb="" moroccon="" clade="" (mp="" 92/93;="" ml="" 99/99;="" bi="" 1="" .="" qhov="" seem="" ntxiv="" ntawm="" c.="" lutea="" (tag="" nrho="" los="" ntawm="" sab="" hnub="" poob="" africa)="" ntxiv="" rau="" c.="" brunneri="" (tsuas="" yog="" ib="" qho="" kev="" nkag="" mus="" rau="" hauv)="" tau="" tsim="" ib="" qib="" ntawm="" lub="" hauv="" paus="" ntawm="" clade="" j="" vim="" txoj="" hauj="" lwm="" ntawm="" c.="" lutea="" accession="" los="" ntawm="" niger="" (ed726)="" sab="" nraum="" lub="" clade="" (mp="" 94/60,="" ml="" 95/69;="" bi="" 1.00/0.99)="" tsim="" los="" ntawm="" lwm="" yam="" accessions.="" qhov="" kev="" txiav="" txim="" siab="" ntawm="" qib="" c.="" lutea="" no="" yuav="" yog="" vim="" qhov="" tsis="" sib="" xws="" ntawm="" cov="" ntaub="" ntawv="" plastid="" thiab="" its.="" tshwj="" xeeb,="" los="" ntawm="" cov="" ntaub="" ntawv="" plastid,="" cov="" c.="" lutea="" accessions="" ntxiv="" rau="" c.="" brunneri="" tau="" suav="" tias="" yog="" ze="" ze="" rau="" c.="" violacea="" thiab="" lwm="" yam="" c.="" lutea="" accessions="" (supplementary="" fig.="" s1),="" piv="" txwv="" li,="" raws="" li="" nyob="" rau="" hauv="" cov="" ntaub="" ntawv="" ua="" ke,="" whereas="" its="" cov="" ntaub="" ntawv="" tso="" rau="" cov.="" ,="" nyob="" rau="" hauv="" ib="" txoj="" hauj="" lwm="" tsis="" daws="" teeb="" meem,="" nyob="" rau="" hauv="" ib="" lub="" clade="" nrog="" c.="" senegalensis="" thiab="" c.="" tubulosa="" rau="" kev="" cais="" tawm="" ntawm="" lwm="" cov="" tswv="" cuab="" ntawm="" clade="" j="" (supplementary="" fig.="">
4. Kev sib tham
Nyob rau hauv qhov tseeb monograph ntawm genusCistanche, Beck-Mannagetta (1930) qhov txawv ntawm plaub pawg (raws li cov seem taxonomic) txawv ntawm calyx nta thiab tus naj npawb ntawm bracteoles. Nrog rau kev zam ntawm monospecific C. sect. Khoom (East Asian Clade), tsis muaj ib qho ntawm Beck Mannagetta cov ntu tau txais kev txhawb nqa raws li monophyletic. Hloov chaw, hom C. sect.Cistanchethiab C. sect. Heterocalyx yog intermixed (nyob rau sab hnub poob Asian Clade thiab dav Clade) thiab ib hom ntawm C. sect. Cistanchella yog paraphyletic thiab zes hauv SouthwestAsian Clade. Ib tug thib plaub loj kab (Northwest African Clade), sawv cev ntawm no los ntawm ib hom tseem yuav piav qhia uas yog morphologically ze rau C. mauritanica, ib txwm muab tso rau hauv C. sect.Cistanchetsis tau raug txheeb xyuas nyob rau hauv ib qho ntawm cov kev faib tawm yav dhau los
Plaub yam kab mob loj uas tau txheeb pom ntawm no tuaj yeem ua tus yam ntxwv morphologically. Tshwj xeeb, ib hom tsiaj ntawm East Asian Clade, C. Sinensis, yog tib yamCistanchehom tsiaj uas muaj qhov sib sib zog nqus quadripartite calyx, hos lwm tus muaj (tsawg kawg) tsib feem ntau connate calyx lobes (Fig. 2). Cov hom kab mob thoob plaws ntiaj teb muaj cov stems glabrous, bracts, thiab calyces, whereas cov neeg ntawm Southwest Asian thiab Northwest African Clade yog tsawg kawg ib nrab plaub hau (woolly los yog lanuginose piv rau arachnoid-lanuginose, feem) (Fig. 2). Thaum kawg, cov tswv cuab ntawm Northwest African Clade muaj qhov ntom ntom arachnoid-lanuginose indumentum thiab dav rhomboid bracts tsis pom nyob rau hauv Southwest Asian Clade (Fig. 2). Yav dhau los hais txog cov cim, piv txwv li, tus naj npawb ntawm bracteoles (ib lossis tsis muaj nyob hauv C. sect.Cistanchellapiv rau ob nyob rau hauv tag nrho lwm seem) thiab cov duab ntawm lub calyx lobes (ntau los yog tsawg sib sib zog nqus lobes thiab tsis sib npaug lobes nyob rau hauv C. sect. heterocalyx piv rau ntiav ntiav thiab sib npaug lobes nyob rau hauv C. sect.Cistanche), tsis txhob xav txog kev sib sib zog nqus sib sib zog nqus thiab, tsawg kawg yog nyob rau hauv cov ntaub ntawv ntawm cov duab ntawm calyx lobes, zoo li tau hloov zuj zus tsawg kawg ob zaug ntawm nws tus kheej.
Ob peb hom tam sim no tsis tau txais kev txhawb nqa raws li monophyletic (Fig. 1), uas tsis txwv rau taxa paub tias nyuaj taxonomically, xws li C. lutea lossis C. tubulosa, tab sis kuj rau taxa suav hais tias yog taxonomically unproblematic, xws li C. . ridgewayana sl orCistanchefissa sl (ntawm no sawv cev los ntawm C. aff. fissa 1 thiab 2). Qhov tsis sib xws ntawm taxonomic thiab phylogenetic units tej zaum yuav yog vim muaj ntau yam, tsis yog kev sib koom ua ke, nrog rau hom misidentification, tsis muaj taxonomic lees paub ntawm morphologically thiab/los yog thaj chaw sib txawv, hybridization, thiab ntiav divergence (hloov hluav taws xob sai). Hom misidentification yog qhov ua rau txoj haujlwm ntawm ob qhov kev nkag ntawm C. tubulosa (GB1 thiab GB2) zes hauv C. laxiflora (Fig. 1). Cov vouchers ntawm cov accessions no tsis muaj rau peb los kho dua, tab sis, raws li txiav txim los ntawm lub vev xaib duab, cov nroj tsuag los ntawm cov cheeb tsam uas nyob ib sab hauv cov suab puam hauv nruab nrab ntawm Tuam Tshoj morphologically pom zoo nrog C. laxiflora subsp. laxiflora (dawb corolla hlab nrog lub teeb liab doog lobes es tsis txhob daj ntseg mus tob daj hauv C. tubulosa). Cov hom no tsis tau tshaj tawm los ntawm Tuam Tshoj, qhov twg cov nroj tsuag hais txog cov taxon no yog teev nyob rau hauv C. tubulosa (Zhang, 1990) los yog, tsis ntev los no, nyob rau hauv C. mongolica (Zhang and Tzvelev, 1998). Tsis muaj taxonomic lees paub ntawm morphologically thiab/los yog geographically sib txawv kab zoo li siv ob qho tib si rau paraphyletic hom, xws li C. ridgewayana sl thiab C. aff. fissa, nrog rau cov hom monophyletic nrog cov qauv phylogenetic muaj zog, xws li C. Flava. Nyob rau hauv txhua rooj plaub no, phylogenetically circumscribed pab pawg sib txawv morphologically thiab thaj chaw thiab, yog li ntawd, tsim nyog taxonomic lees paub (cov ncauj lus kom ntxaws taxonomic kev kho mob yuav muab nyob rau hauv ib qho kev tshaj tawm tom ntej). Ob leeg, hom misidentification thiab tsis txaus taxonomic daws yog teeb meem tshwm sim nyob rau hauv holoparasitic Orobanchaceae, tshwj xeeb tshaj yog nyob rau hauv Orobanche thiab lwm yam genera (Manen li al., 2004; Schneeweiss li al., 2004a; Schneeweiss li al., 2009; 2 Schneeweiss, 2009;
Hybridization yog ib qho tshwm sim tshwm sim hauv cov paj ntoo (Rieseberg thiab Carney, 1998; Payseur thiab Rieseberg, 2016) thiab kuj tau tshaj tawm rauCistanche. Tshwj xeeb, Beck-Mannagetta (1930) tau piav qhia txog kev sib xyaw ntawm C. lutea (raws li C. tinctoria f. lutea) thiab C. violacea as nothospecies, C. hybrid. Cov hybrids no tej zaum yuav yog morphologically nruab nrab, ib yam li cov ntaub ntawv rau accession ED686, los yog tej zaum yuav zoo li ib tug ntawm cov niam txiv, ib yam li cov ntaub ntawv rau accession ED1012; qhov kev nkag mus tom kawg no tau raug txiav txim siab thaum xub thawj ua C. violacea, tab sis ntawm kev tshuaj xyuas rov qab, provoked los ntawm nws txoj hauj lwm phylogenetic nyob rau hauv C. lutea sl, tig tawm los ua ib tug hybrid thiab. Piv txwv li niam txiv qub txeeg qub teg ntawm plastid genome (raws li qhia rau Orobanchaceae Rhinanthus angustifolius: Vrancken thiab Wesselingh, 2010) hauvCistanche, lub xub ntiag ntawm leej niam ITS ribotype nyob rau hauv hybrid accession ED686 (nws tsis muaj nyob rau accession ED1012) yuav zoo ib yam nrog cov tsim ntawm tiam tom ntej hybrids thiab / los yog backcrosses, ua rau plastid capture. Kev ntes Plastid kuj tseem tuaj yeem piav qhia qhov tsis sib xws ntawm cov cim nuclear thiab plastid. Piv txwv li, C. aff. tubulosa los ntawm Arabian ceg av qab teb, morphologically txawv nyob rau hauv ob peb yam ntxwv los ntawm thoob plaws C. tubulosa, tsim ib tug clade nyob ze rau C. senegalensis nyob rau hauv cov ntaub ntawv ITS tab sis tau muab cais ua ob sib txawv clades nyob rau hauv lub plastid thiab cov ntaub ntawv sib xyaw ua ke nrog C. senegalensis. (clade H) lossis kev nkag ntawm C. tubulosa subsp. tubulosa (clade I; ob qho tib si C. senegalensis thiab C. tubulosa subsp. tubulosa kuj tshwm sim ntawm Arabian Peninsula).
Ib qho kev hloov rau introgression rau piav qhia qhov tsis sib xws ntawm phylogenetic txoj hauj lwm inferred los ntawm nuclear piv rau plastid cov ntaub ntawv yog cov kab lus tsis tiav, uas yuav cuam tshuam rau nuclear thiab plastid genomes sib txawv vim qhov sib txawv ntawm lawv cov pej xeem qhov ntau thiab tsawg (hauv plastid genomes tsuas yog ib nrab loj li hauv nuclear genomes hauv ib pawg monoecious xws liCistanche). Kev txheeb xyuas tsis tiav yuav tsum yog qhov tshwj xeeb tshaj yog nyob rau hauv cov ntaub ntawv ntawm qhov sib txawv me me thiab hluav taws xob nrawm (Maddison thiab Knowles, 2006), uas pom tau tias yog qhov xwm txheej hauv Widespread Clade, qhov twg cov clade suav nrog Clades G rau J yog cov ceg luv luv. (Daim duab 1). Kev txheeb xyuas tsis tiav tej zaum yuav yog lub luag haujlwm rau qhov tsis muaj qhov sib txawv ntawm cov molecular thiab / lossis kev sib koom ua ke ntawm cov kab mob sib txawv hauv C. lutea sl (ie, C. aff. lutea 1 thiab C. aff. lutea 2). Disentangling incomplete lineage sorting from introgression will need coalescent-based method (Blanco-Pastor li al., 2012) zoo tshaj plaws nrog rau cov ntaub ntawv ntawm cov ntaub ntawv nuclear (Bravo li al., 2019), uas nyob rau hauv qhov tsis muaj cov khoom taxonomic moj khaum thiab txaus cov ntaub ntawv tsis txaus. ua tau.
Tus tswv tsev tshwj xeeb yog ib qho tseem ceeb evolutionary quab yuam nyob rau hauv holoparasitic Orobanchaceae (Schneider li al., 2016).CistancheTxawm li cas los xij, zoo li yog qhov tshwj xeeb, vim tias muaj ntau hom tsiaj tau pom nyob hauv cov tswv cuab ntawm ntau tsev neeg cog, feem ntau Amaranthaceae (xws li Chenopodiaceae) thiab Polygonaceae, tab sis kuj Fabaceae, Zygophyllaceae, Tamaricaceae, Rosaceae, Nitrariaceae, thiab Salvadoraceae. Raws li nyob rau hauv Orobanche (Manen li al., 2004), tib tus tswv tsev yuav raug muab faib los ntawm ntau yam tsis cuam tshuam.Cistanchehom (piv txwv li, Haloxylon ammodendron yog tus tswv tsev rau C. deserticola los ntawm Southwest Asian Clade thiab rau C. phelypaea los ntawm Widespread Clade). Cov xwm txheej ntawm cov tswv tsev tshwj xeeb (piv txwv li, C. senegalensis feem ntau ntawm Acacia / Fabaceae; feem ntau yog Fabaceae ua tswv rau C. Flava subsp. nov. piv rau feem ntau Calligonum bungei / Polygonaceae rau C. Flava subsp. Flava) tuaj yeem cuam tshuam txog kev nplua nuj ntawm cov tswv tsev tsim nyog. hom es tsis yog tus tswv tshwj xeeb tiag tiag. Kev ntsuam xyuas txhua lub luag haujlwm ntawm tus tswv tshwj xeeb rau kev sib txawv ntawm Cistanche yuav xav tau kev daws teeb meem zoo ntawm hom phylogeny thiab cov ncauj lus kom ntxaws tswv yim txaus.5. Cov lus xaus
Qhov no yog thawj qhov kev tshuaj ntsuam xyuas molecular phylogenetic ntawm cov nroj tsuag holoparasitic.Cistanche, dav faib nyob rau hauv arid regions ntawm lub qub ntiaj teb no. Plaub loj clades nyob rau hauvCistanchetau raug txheeb xyuas tias tsuas yog ib feem sib raug rau cov kab ke uas tau lees paub (Beck Mannagetta, 1930) thiab feem ntau qhia txog thaj chaw muaj zog. Whereas Southwest Asian Clade nthuav tawm cov qauv phylogenetic muaj zog ntawm thiab ib nrab nyob rau hauv cov tsiaj (East Asian thiab Northwest African Clade yog monospecific), phylogenetic kev daws teeb meem nyob rau hauv Dav Clade feem ntau qis thiab cuam tshuam los ntawm qhov tsis sib xws ntawm nuclear thiab plastid cov cim, uas tsawg kawg yog ib feem. yog vim hybridization/introgression thiab/los yog tsis tiav lineage sorting. Molecular phylogenetic pov thawj thiab cov txiaj ntsig ntawm morphological rov ntsuam xyuas ntawmCistanchehom ntawv qhia tias hom kev sib txawv hauvCistancheyog tam sim no underestimated (saib taxa qhia raws li sp. nov. los yog subsp. nov. nyob rau hauv cov ntawv nyeem). Txawm hais tias qee hom tseem yuav tsum tau muab tso rau hauv ib qho kev tshawb fawb molecular phylogenetic (piv txwv li, C. mauritanica) thiab cov cim ntxiv yuav xav tau los daws txhua hom kev sib raug zoo, lub suab phylogenetic hypotheses tau nthuav tawm ntawm no muab lub hauv paus tseem ceeb rau kev txuas mus ntxiv cytogenetic, taxonomic, thiab biogeographic. kev tshawb fawb nyob rau hauv no genus.
cistanche bienfaits
Nyiaj txiag
Txoj haujlwm no tau txais kev txhawb nqa ib nrab los ntawm SYNTHESYS nyiaj txiag los ntawm European Community Research Infrastructure Action nyob rau hauv FP7 "Capacities" Program ntawm Real Jardín Botánico (ES-TAF-1663). Peb xav ua tsaug rau University of Bonn, OeAD (Österreichische Austauschdienst) thiab DAAD (Deutscher Akademischer Austauschdienst) rau kev txhawb nqa nyiaj txiag.
CREDiT daim ntawv qhia txog kev sau ntawv
Najibeh Ataei: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Project Administration, Visualization, Sau - thawj daim ntawv, Sau - tshuaj xyuas & kho.Gerald M. Schneeweiss: Conceptualization, Supervision, Sau -review & editing. Miguel Angel García: Cov peev txheej. Michael Krug: Kev txheeb xyuas raug cai. Marcus Lehnert: Sau - thawj tsab ntawv. JafarValizadeh: Cov peev txheej. Dietmar Quandt: Conceptualization, Formal analysis, Project Administration, Resources, Supervision, Sau - tshuaj xyuas & kho
Kev lees paub
Ua tsaug rau Edit Korpinos, uas tau muab kev pab ntawm Jodrell lab hauv Kew thiab rau cov kws kho mob herbarium (TARI, BM, IRAN, BONN, USB, P, TUH, E, W, KAS, MSB, K, B, G, PEY , M, UG, BR, GUH, MA, WU) rau kev xa nyiaj qiv thiab duab. Peb ua tsaug Susann Wicke rau primer tsim thiab lab kev pab txhawb nqa. Peb txaus siab rau Dr. Hossein Akhani ntawm TehranUniversity thiab Mrs. Robabeh Shahi Shavvon ntawm Gilan University tau muab qee cov khoom siv DNA los ntawm Iran. Peb ua tsaug rau Dr. FedericoLuebert thiab Juliana Chacon ntawm Nees lub koom haum rau lawv cov lus pab.
Cov ntaub ntawv:
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