First Nuclear DNA Amounts in more than 300 Angiosperms

16 Pages·2005·0.16 MB·English

Annalsof Botany96:229–244,2005
doi:10.1093/aob/mci170,available onlineatwww.aob.oupjournals.org
First Nuclear DNA Amounts in more than 300 Angiosperms
B. J. M. ZONNEVELD1, I. J. LEITCH2 and M. D. BENNETT2,*
1Institute ofMolecular Plant Sciences, LeidenUniversity, Leiden, The Netherlandsand 2Jodrell Laboratory,
Royal Botanic Gardens, Kew, Richmond, SurreyTW9 3AB,UK
Received:3February2005 Returnedforrevision:30March2005 Accepted:10April2005 Publishedelectronically:19May2005
D
o
(cid:1)BackgroundandAimsGenomesize(DNAC-value)dataarekeybiodiversitycharactersoffundamentalsigni- w
n
ficanceusedinawidevarietyofbiologicalfields.Since1976,Bennettandcolleagueshavemadescatteredpublished lo
a
and unpublished genome size data more widely accessible by assembling them into user-friendly compilations. d
e
Initiallythesewerepublishedashardcopylists,butsince1997theyhavealsobeenmadeavailableelectronically d
(see the Plant DNA C-values database www.kew.org/cval/homepage.html). Nevertheless, at the Second Plant fro
GenomeSizeMeetingin2003,Bennettnotedthatasmanyas1000DNAC-valueestimateswerestillunpublished m
andhenceunavailable.Scientistswerestronglyencouragedtocommunicatesuchunpublisheddata.Thepresent h
workcombinesthedatabasingexperienceoftheKew-basedauthorswiththeunpublishedC-valuesproducedby ttp
s
Zonneveldtomakealargebodyofvaluablegenomesizedataavailabletothescientificcommunity. ://a
(cid:1)MethodsC-valuesforangiospermspecies,selectedprimarilyfortheirhorticulturalinterest,wereestimatedbyflow c
a
cytometryusingthefluorochromepropidiumiodide.Thedatawerecompiledintoatablewhoseformissimilarto d
e
previouslypublishedlistsofDNAamountsbyBennettandcolleagues. m
(cid:1)KeyResultsandConclusionsThepresentworkcontainsC-valuesfor411taxaincludingfirstvaluesfor308species ic
.o
not listed previously by Bennett and colleagues. Based on a recent estimate of the global published output of u
p
angiospermDNAC-valuedata(i.e.200firstC-valueestimatesperannum)thepresentworkequals1(cid:2)5yearsof .c
averageglobalpublishedoutput;andconstitutesover12%ofthelatest5-yearglobaltargetsetbytheSecondPlant om
Genome Size Workshop (see www.kew.org/cval/workshopreport.html). Hopefully, the present example will /a
encourage others to unveil further valuable data which otherwise may lie forever unpublished and unavailable ob
forcomparativeanalyses. /a
rtic
Key words: Genome size, C-value, plant DNA C-values database, Genome Size Initiative (GSI), Plant Genome Size le
workshopandDiscussionMeeting,angiosperm,monocots. -ab
s
tra
c
INTRODUCTION (e.g. Knight et al., 2005; Leitch et al., 2005; Price et al., t/9
6
2005) and are the main source of information on plant /2
TheDNAamountintheunreplicatedgameticnuclearchro- genomesizescitedintheliterature.Publishedlistsofangio- /22
mosome complement (1C-value) is highly characteristic 9
sperm DNA amounts have been cited over 1500 times, /2
for taxa and varies over 1000-fold between plant species. 9
whilsttheelectronicdatabasehasreceivedover60000hits 9
Species’ DNA amounts [C-value and genome size (1Cx)] 1
since 2001. 62
are key diversity characters of fundamental significance Bennett et al. (2000) and Bennett and Leitch (2005a) by
with many important uses (Bennett and Leitch, 2005a). notedthatC-valuesforprobablyasurprisinglylargenumber gu
However, it is often difficult to know if a genome size e
ofplantspecieshadbeenestimatedbut,forvariousreasons, s
measurementexistsforataxon,orwheretofindit,assuch notpublishedandhencewereunavailable.Indeed,thiswas t on
valuesarewidelyscatteredintheliteratureorunpublished. suspected to apply to hundreds of species overall whose 15
Csppuoebnrlmsieshqseupdeenc(teile.ygs,,.sBcionemcnenpei1lt9etd7a6nf,dolrSisrmtesfietohrfe,nD1c9Ne7A6p)u,arampnoodsuetnsh,teshdianavteaanbhgeaievone- ginefnoormmeatiosinzesoftocgoenthsiedrercaobnlestitpuotteenatinalunvtaalpupee.dScpioeonltisotsf April 2
beenpooledandreleasedinelectronicformsince1997(e.g. have therefore been encouraged to communicate them for 019
inclusioninthePlantDNAC-valuesdatabase,thusmaking
the Angiosperm DNA C-values database). To extend the
them accessible for comparative studies.Thepresent work
coveragetoincludeotherplantgroups,databasesforgym-
isproofofthishypothesisandprovidesamodelexampleof
nosperms,pteridophytesandbryophyteswereaddedin2001
collaborationtotransferalargebodyofpreviouslyinvisible
tocreatethePlantDNAC-valuesdatabase(release1(cid:2)0)with
plant genome size data firmly into the public domain.
C-valuesfor3864landplants.InDecember2004thePlant
Twoofthepresentauthorshaveworkedtogetherforthe
DNA C-values database was further updated (release 3(cid:2)0)
pastdecadetoproducepooledlistsofplantDNAC-values
and included, as a novel feature, C-values for 253 algal
for reference purposes in both hard copy (Bennett and
species. Release 3(cid:2)0 has DNA amounts for almost 5000
Leitch, 1995, 1997, 2005a; Bennett et al., 2000) and elec-
species from over 500 original sources, including first val-
tronicversions(http://www.kew.org/genomesize/homepage).
ues for 628 angiosperms not previously included (http://
Meanwhile the present first author has made genome size
www.kew.org/genomesize/homepage).Theselistsanddata-
estimates for angiosperm taxa in several genera, including
bases have been widely used for comparative studies
Helleborus (Zonneveld, 2001), Hosta (Zonneveld and
*[email protected] VanIren,2000,2001), Galanthus(Zonneveldet al.,2003)
ªThe Author2005. PublishedbyOxfordUniversityPressonbehalf ofthe Annalsof BotanyCompany. Allrightsreserved.
For Permissions, pleaseemail: [email protected]
230 Zonneveld et al. — Nuclear DNA Amounts in Angiosperms
and Agapanthus (Zonneveld and Duncan, 2003), some TABLE 1. Species used as calibration standards to estimate
ofwhichhavesubsequentlybeenincludedinarecentcom- nuclear DNA C-values
pilationbyBennettandLeitch(2005a).Whilecorrespond-
ing with Bennett and Leitch about these data and planned Assumed Abbreviationused
futurepublications,Zonneveldnotedthathehadalsomade 2CDNA incolumn13of
Species amount(pg) theAppendix
C-value estimates for taxa in various genera that seemed
most unlikely to be published. It emerged that this sample Agavestricta 7.8 A.stricta
wassubstantialandincludedfirstestimatesforhundredsof Hordeumvulgare‘Sultan’* 10.0 Hordeum
miscellaneous species, measured mostly out of curiosity. Agaveamericana 15.9 A.americana
Giventheirpotentialvalue,itwasdecidedtocombinethe Agavesisalana 20.2 A.sisalana
Cliviaminiata 39.0 Clivia
taxonomic and databasing experience of the Kew-based
authors with information amassed in The Netherlands to *The2Cvalueof10pgforHordeumvulgare‘Sultan’wasobtainedusing D
produce a further reference list in a form similar to other Agaveamericana(2C=15(cid:2)9pg)asthecalibrationstandard(seeentrynumber ow
recentplantC-valuereferencelists(e.g.BennettandLeitch, 161qintheAppendix).ThisvalueisslightlylowerthanthatgiveninBennett nlo
2005a; Kapraun, 2005). The present work is the product andSmith(1991)of2C=11(cid:2)12pg.IntheAppendixH.vulgarewasusedasa ad
of that collaboration. Analysis of the completed Appendix calibrationstandardforonlyonespeciesCatharanthusroseus(2C=1(cid:2)4pg; ed
table shows that it contains C-values for 411 angiosperm seeentrynumber82binAppendix).IfthehighervalueforH.vulgaregiven fro
byBennettandSmith(1991)isusedthenthe2CvalueofC.roseuswould m
taxa,includingfirstvaluesfor308speciesnotlistedinany be1(cid:2)6pg. h
previous compilation by Bennett and colleagues. ttp
s
flowcytometeravailableforuse,asthepeakstendedtobe ://a
c
lost in the debris fluorescence. a
d
MATERIALS AND METHODS e
m
ic
Plantmaterials .o
RESULTS AND DISCUSSION up
The present work examined angiosperm taxa available as .c
o
seed or plants growing in The Netherlands selected prim- ComparisonofC-valuedataintheAppendixtablewiththose m
arily for their horticultural interest. The Netherlands has a previouslyreported /ao
b
ltroaexandagifltyoraradhviotairiotlinacbuolleftufgrorero,wrceoisnnegsaeraqcnuhdepnbutlrryepeosdusiecnshg.pmlaanntymaantgerioiaslpsearrme Tcoom(1ap)sasCeris-svsoantlhsueewsqeufroaerlimt1y0a3odfes.pCe-cviaelsueindtahteaAinpptheendAixppwenerdeixcotwmo- /article-a
b
EstimationofDNAC-values pthaereAdpwpeitnhdi‘xp’r)imCe-v’a(lsueees nfoorteth(ea)saimne‘Nspoetecsiesonlisctoedmpinilitnhge strac
DNA 2C-values were estimated using flow cytometry Plant DNA C-values database (Bennett and Leitch, 2004). t/96
with the fluorochrome propidium iodide (PI) as described Figure1Ashowsthatgenerallythetwodatasetswereingood /2/2
byZonneveldandVanIren(2001).Briefly,somaticnuclei agreement (R2 = 0(cid:2)9068). However, there were 18 species 29
wereisolatedfromleavesbyco-choppingthemwithoneof where previously published ‘prime’ C-values differed by /29
9
the internal calibration standards (Table 1) in nuclei isola- >30% from those given in the Appendix. These species 1
6
tion buffer. After adding 2 mL PI solution (50 mg PI/l in are represented by open circles in Fig. 1A. The cause(s) 2 b
half-strength isolation buffer) the suspension was filtered. of the discrepancies was not clear but possibilities include y g
Nucleifluorescencewasmeasured30minaftertheaddition taxonomicerrorsorgenuineintraspecificvariation.Forsix ue
s
of PI using a Partec CA-II flow cytometer. In most cases, ofthespeciesthedifferentC-valuesreportedweremultiples t o
severalsamples,eachwithatleast5000nuclei,weremeas- of previous estimates, suggesting that polyploidy may be n 1
ured.The2C-valueofthesamplewascalculatedasfollows: responsible. Nevertheless, as no chromosome counts were 5 A
(ammeoaunnotf(spagm)polfetpheeaksta(cid:3)ndmaredanspoefcsiteasnd(aserdepTeaabkl)e·12)C. ODtNheAr mmaacdreofpohrylalnay),stpheecrieosleinofthpeolAypplpoeinddyixis(uenxccoepntfirHmyeddr.angea pril 2
0
peaks corresponding to higher C-values (i.e. 4C, 8C and (2) A comparison was made between the C-values for 19
16C,etc.)weresometimesobservedbuttheirDNAamounts seven species in the Appendix and the C-values for these
were not calculated. calibration standards given in Bennett and Smith (1991).
Thesetwodatasetsshowedverycloseagreement(Fig.1B)
Internalcalibrationstandards (R2=0(cid:2)9965).Overall,itwasconcludedthattheAppendix
C-value data are of high quality.
Five calibration standards, differing in DNA amount,
were used (Table 1). In most cases Agave americana was
RepresentationofC-valuedatainAppendix
chosen, as previous work showed it to be a reliable and
reproducible standard, giving clean peaks with low CVs The present Appendix gives C-values for 411 species,
(Zonneveld and Van Iren, 2001). If nuclear DNA content of which 308 (75%) are for species not included in any
of a test species overlapped with that of A. americana, previous compilation by Bennett and colleagues (Bennett
another calibration standard was used whose C-value fell andSmith,1976,1991;Bennettetal.,1982,2000;Bennett
closetothatoftheunknown.Specieswith2C-valuesbelow and Leitch, 1995, 1997, 2005a)or listed in the Plant DNA
2(cid:2)0pgcouldusuallynotbemeasuredwiththePartecCA-II C-values database (release 3(cid:2)0, December 2004). Bennett
Zonneveld et al. — Nuclear DNA Amounts in Angiosperms 231
60 y = 0·8582x + 0·8914 20 y = 0·957x + 0·5172
g) A R2 = 0·9008 pg) B R2 = 0·9965 Triticum aestivum
ported (p 40 d Smith ( 15 Vicia fabAallium cepa
usly re nett an 10
o n
evi Be Secale cereale
pr 20 y
value alue b 5 PisHuomr dseautimvu vmulgare
1C- 1C-v Senecio vulgaris Dow
n
0 0 lo
0 20 40 60 0 5 10 15 20 a
d
e
1C-value in Appendix (pg) 1C-value in Appendix (pg) d
fro
FIG. 1. ComparisonsbetweenC-valuesforspecieslistedintheAppendixand‘prime’C-valuesforthesamespecieslistedinthePlantDNAC-values m
database(release3(cid:2)0,December2004).(A)Comparisonforall103species.Opencirclescorrespondtothe18specieswhoseC-valuesintheAppendixdiffer h
by>30%fromthe‘prime’C-valuesinthedatabase.(B)ComparisonbetweenC-valuesforsevenspecieslistedintheAppendixandtheacceptedC-valuesfor ttp
s
thesecalibrationstandardsgiveninBennettandSmith(1991). ://a
c
a
d
and Leitch (2005a) analysed the global output of DNA TABLE 2. RepresentationofC-valuedataintheAppendixat em
C-values for angiosperms during the past half century. various taxonomic levels ic
.o
They concluded that the 1997 Angiosperm Genome Size u
p
Workshophadstimulatedalargeincreasefromanaverage No.listedin No.offirst .c
o
ofabout100perannumintheearly1990stoalmost200per Appendixtable C-values m
/a
annum for 1998–2002. The total number of first C-values o
Species 411 308 b
forspeciesinthepresentworkthereforeequals1(cid:2)5yearsof /a
average global published output at the record levels BMaosnaolcaontgsiosperm 2583 1956 rticle
achievedinrecentyears.Byanymeasure,itisasignificant Eudicots 150 107 -a
contribution to knowledge of this important character. Genera 187 85 bs
Indeed the present data sample (all from one source) Basalangiosperm 6 2 tra
increases the total sample of angiosperm species with MEuodniococotsts 16147 5267 ct/96
genome size estimates by about 7(cid:2)4%. Moreover, it con- Family 67 1 /2
stitutes over 12% of the global target set by the Second Basalangiosperm 5 0 /22
Plant Genome Size Workshop (namely to estimate first Monocots 21 0 9/2
Eudicots 41 1 9
DNA C-values for about 2500 angiosperm species within 9
1
a quinquennium; see http://www.rbgkew.org.uk/cval/ 62
workshopreport.html). Clearly, one worker can still make by
a significant contribution to the global pool of information concluded that significant progress to improve familial gu
e
on genome sizes. Failing to make such information visible representationofangiospermfamilieswasunlikelywithout s
could seriously impair the scope of the available database careful targeting and the present data for a large sample t on
and limit the scope of comparative studies. of untargeted taxa confirm this expectation. Thus, while 15
Table2showshowmanyoftheC-valuesintheAppendix the Appendix includes 308 first C-values for species, Ap
are first values at the species, generic and family level for only one of these is also the first value for a family (i.e. ril 2
thethreemajorangiospermgroups(i.e.basalangiosperms, Tamaricaceae). 01
9
monocotsandeudicots).Althoughmonocotsconstituteonly Also as expected, given the lack of specific targeting,
approx. 20% of all angiosperm species, most of the new the present sample does not make major contributions to
C-valuesintheAppendixareformonocots(i.e.64and65% filling gaps for any other groups identified as significantly
of first C-values are for monocot species and genera, under-represented in the database (e.g. for taxa from
respectively). bog, fen, tundra, alpine and desert environments, or for
Nevertheless, while the present data make a significant halophytic, insectivorous, parasitic, saprophytic and endo-
contributiontoknowledgeofC-valuesatthespecieslevel, phytic species; Bennett and Leitch, 2005a). Nevertheless,
therehasbeennosignificantimprovementinrepresentation thepresentsampledoesincludefirstvaluesforafewtaxaof
at the family level. The 1997 Angiosperm Genome Size such interest including two parasitic species Orobanche
Workshop recommended a goal of complete familial rep- hederae and Rhinanthus glabra (both in Orobanchaceae)
resentation by 2002, as a first C-value was available for with 1C values of 2(cid:2)65 and 3(cid:2)05pg, respectively. The
only about 30% of families. Bennett et al. (2000) noted table also includes two desert species Opuntia microdasys
thatinasixthsupplementarylistofC-valuesfor691species (1C = 2(cid:2)24pg) and Rebutia albiflora (1C = 1(cid:2)91pg), both
only 12 were also first estimates for families. Later they in the Cactaceae, and a number of succulents from the
232 Zonneveld et al. — Nuclear DNA Amounts in Angiosperms
TABLE 3. Theminimum,maximum,mean,median,mode,and Futureplans
rangeof1CDNAvaluesfor411speciesintheAppendixcom-
Participants at the 2003 Plant Genome Size Workshop
paredwiththecorrespondingstatisticsfor4119speciesinthe
formalized their international collaboration as the Genome
Plant DNA C-values database(release 3.0,December 2004)
SizeInitiative(GSI;BennettandLeitch,2005b),whoseaims
aretoimprovetheavailability,qualityandunderstandingof
PlantDNAC-values
database(release3.0, genome size data, and to provide a focus for monitoring
Appendix December2004) progress and facilitating discussions in a holistic genomic
context.Further,inakeynoteaddressopeningthe2003Plant
Minimum1C-value(pg) 0.6 0.1 Genome Size Discussion Meeting, Bennett reiterated the
Maximum1C-value(pg) 95.0 127.4 potential value of unpublished genome size data, and the
Mean1C-value(pg) 11.7 6.1
needtomakeitvisibleasfollows:‘Thereareprobablyover
Median1C-value(pg) 6.6 2.7
Mode1C-value(pg) 5.0 0.6 1000 prime C-value estimates for plant species, equal to Do
Range(max/min) 94.4 1274.0 4 years of known global output, unlocated in publications wn
orunpublished.Suchgenomesizedataarealmostvalueless loa
d
ifpotentialuserscannoteasilyaccessandassessthem.Itis e
d
families Asparagaceae (Agave, Ledebouria), Bromeliaceae impossibletoover-emphasisetheimportanceofpublishing fro
C-value data or linking them to the database by informing m
S(Teidlulamn)d,sia)A,poCcyransascuelaaeceae(Ce(rAoepoengiiuam, , HGoroadpiato,petHalouyma,, theco-ordinatorsofitsexistence,sothatotherscanassessits http
qualityanduseit’.Thepresentworksupportsthissentiment s
Catharanthus, Stapelia), Xanthorrhoeaceae (Aloe) and andtheaimsofGSI.Hopefully,itwillencourageothersto ://a
Asteraceae (Senecio) in which the C-values ranged from ca
emulate this example and unveil further useful data which d
1(cid:2)11 to 16(cid:2)85pg. e
otherwise may lie forever unpublished. m
ic
.o
u
TherangeofC-valuesintheAppendixcompared p
withexistingdata LITERATURE CITED .co
m
APGII.2003.AnupdateoftheAngiospermPhylogenyGroupclassification /a
Table 3 compares the minimum, maximum, mean, o
fortheordersandfamiliesoffloweringplants.BotanicalJournalofthe b
median, mode and range of DNA amounts for all 411 spe- LinneanSociety141:399–436. /a
cies listed in the Appendix, with the 4119 species listed in Bennett MD, Leitch IJ. 1995. Nuclear DNA amounts in angiosperms. rtic
the Plant DNA C-values database (release 3(cid:2)0, December AnnalsofBotany76:113–176. le-a
BennettMD,LeitchIJ.1997.NuclearDNAamountsinangiosperms:583 b
2004). s
The range of C-values in the present sample (1C = 0(cid:2)6– newestimates.AnnalsofBotany80:169–196. tra
BennettMD,LeitchIJ.2004.PlantDNAC-valuesdatabase(release3(cid:2)0, c
95(cid:2)5pg) falls within that reported for the Plant DNA December2004).http://www.kew.org/cval/homepage.html t/9
6
C-valuesdatabase(1C=0(cid:2)1–127(cid:2)4pg),althoughitissome- BennettMD,LeitchIJ.2005a.NuclearDNAamountsinangiosperms: /2
what enriched in species with larger genomes, leading to progress,problemsandprospects.AnnalsofBotany95:45–90. /22
BennettMD,LeitchIJ.2005b.Plantgenomesizeresearch:afieldinfocus. 9
higher mean and modal values compared with the larger AnnalsofBotany95:1–6. /29
sampleinthedatabase(Table3);thustheminimumC-value Bennett MD, Smith JB. 1976. Nuclear DNA amounts in angiosperms. 91
intheAppendixisthesameasthemodalvalueforspecies PhilosophicalTransactions of the Royal Society of London. Series 62
inthedatabase.Thisprobablyreflectsthegreaterproportion B.BiologicalSciences274:227–274. by
of monocots in the present sample with large genomes. Bennett MD, Smith JB. 1991. Nuclear DNA amounts in angiosperms. gu
PhilosophicalTransactions of the Royal Society of London. Series e
Indeed the 43 species (approx. 10%) with the largest gen- s
omes in the Appendix are all monocots from the orders BenneBt.tBMioDlo,SgimcaitlhSJcBie,nHceessl3o3p4-H:3a0r9r–is3o4n5J.S.1982.NuclearDNAamounts t on
Asparagales, Commelinales or Liliales; these groups have inangiosperms.ProceedingsoftheRoyalSocietyofLondon.SeriesB. 15
previously been shown to contain species with the largest BiologicalSciences216:179–199. Ap
recorded angiosperm genomes (Leitch et al., 1998; Soltis BennesptteMrmDs,aBndhtahnediromlPo,dLerenitucsheIsJ:8.20070n0e.wNeusctilmeaartDesN.AAnanmalosuonftBsointaannyg8io6-: ril 2
et al., 2003). (NB The presence of only eight species with 859–909. 01
9
2C-values <2(cid:2)0pg is, in large part, due to technical diffi- BrandhamPE,WestJP.1993.CorrelationbetweennuclearDNAvalues
culties encountered in estimating C-values in species with anddifferingoptimalploidylevelsinNarcissus,HyacinthusandTulipa
cultivars.Genetica90:1–8.
smaller genomes, as noted in Materials and methods.)
Cavalier-SmithT.1985.Theevolutionofgenomesize.Chichester:John
Analysis at the family level showed that in half of the
Wiley&Sons.
66familiesalreadyrepresentedinthePlantDNAC-values DabrowskaJ.1992.Chromosome numberandDNAcontentintaxaof
database, the new data did not increase the range of AchilleaL.inrelationtothedistributionofthegenus.PraceBotaniczne
C-values previously reported for a family. In only eight 49:1–84.
Kapraun DF. 2005. Nuclear DNA content estimates in multicellular
families was the minimum C-value decreased as a result
eukaryoticgreen,redandbrownalgae:phylogeneticconsiderations.
ofnewdata,whereasin25familiesthemaximum C-value AnnalsofBotany95:7–44.
increased, sometimes greatly. For example, in Garryaceae KnightCA,MolinariN,PetrovDA.2005.Thelargegenomeconstraint
and Escalloniaceae, which both previously had a C-value hypothesis:evolution,ecologyandphenotype.AnnalsofBotany95:
177–190.
forjustonespecies,theadditionofanestimateforonenew
Leitch IJ, Chase MW, Bennett MD. 1998. Phylogenetic analysis of
speciesincreasedtherangeofC-valuesforeachfamilyover
DNAC-valuesprovidesevidenceforasmallancestralgenomesize
8- and 15-fold, respectively. infloweringplants.AnnalsofBotany82:85–94.
Zonneveld et al. — Nuclear DNA Amounts in Angiosperms 233
LeitchIJ,SoltisDE,SoltisPS,BennettMD.2005.EvolutionofDNA S.pendulusandS.radicans),however,thedifferentC-values
amounts across land plants (Embryophyta). Annals of Botany 95: so strongly suggested the presence of polyploidy that they
207–217. weregivenseparateentrynumbersinaccordancewithpre-
PriceHJ,DillonSL,HodnettG,RooneyWL,RossL,JohnstonJS.2005.
viouspractice(e.g.BennettandLeitch,2005a).
GenomeevolutioninthegenusSorghum(Poaceae).AnnalsofBotany
95:219–228. For103species,whereoneormoreC-valuehadpreviously
Soltis DE, Soltis PS, Bennett MD, Leitch IJ. 2003. Evolution of been listed by Bennett and colleagues, the estimate in the
genome size in the angiosperms. American Journal of Botany 90: Appendixwasgivenanumberandthenextavailableletterin
1596–1603.
thealphabet.Forexample,threepreviousC-valueshavebeen
United Nations Environment Programme. 1992. Convention on bio-
reported for Allium ampeloprasum, thus the entry for this
logical diversity. Nairobi, Kenya: United Nations Environment
Programme. speciesintheAppendixwasassignedtheletter‘d’.
ZonneveldBJM.2001.NuclearDNAcontentsofallspeciesofHelleborus
(b) Taxonomic authorities. Taxonomic authorities of
(Ranunculaceae)discriminatebetweenspeciesandsectionaldivisions.
PlantSystematicsandEvolution229:125–130. the species studied were taken from the International Do
ZonneveldBJM,DuncanGD.2003.Taxonomicimplicationsofgenome PlantNamesIndex(IPNI)(http://www.ipni.org/index.html). w
n
sizeandpollencolourandvitalityforspeciesofAgapanthusL’Heritier Asuperscript‘b’followingaspeciesnameindicatesthatthe lo
a
(Agapanthaceae).PlantSystematicsandEvolution241:115–123. d
taxonomicauthoritywasunknownorunclear(i.e.morethan e
Zonneveld BJM,VanIren F.2000.FlowcytometricanalysisofDNA d
contentinHostarevealsploidychimeras.Euphytica111:105–110. oneentryinIPNI). fro
ZonneveldBJM,VanIrenF.2001.Genomesizeandpollenviabilityas m
taxonomiccriteria:ApplicationtothegenusHosta.PlantBiology3: (c)Familyandhigherordergroup.Thefamily,orderand h
176–185. highergroupgivenforeachspeciesinAppendixcolumns3, ttps
ZonneveldBJM,GrimshawJM,DavisAP.2003.Thesystematicvalueof 4 and 5, respectively, follows the Angiosperm Phylogeny ://a
nuclearDNAcontentinGalanthus.PlantSystematicsandEvolution Group(APG)system(APGII,2003).Thefollowingabbre- c
a
241:89–102. viationswereused:BA=basalangiosperm;M=monocot; de
m
E=eudicot. ic
APPENDIX .o
(d) Chromosome number, ploidy and voucher informa- up
NotesoncompilingtheAppendix tion.ThesentimentoftheConventiononBiologicalDivers- .co
m
C-valuedatawerecompiledintotheAppendixtablewhose ity(UnitedNationsEnvironmentProgramme,1992),which /a
form is similar to that of previous lists of DNA amounts noted the need to make biodiversity data available despite ob
b1y99B1;enBneetntnaenttdectolalel.a,gu1e9s82(i;.eB. eBnennenttettanadndLSemitcithh,, 11999756,, iimnptheerfAecptipoennsd,imxegreitnsesruapllpyolrat.ckThauvs,owuchhileertahnedspinecfoiermslaistitoend /article
1997; Bennett et al., 2000). Additional information on onchromosomenumberandploidylevel,theycan makea -ab
compilingtheAppendixisgivenbelow. valuablecontributiontoknowledgeofplantC-values. stra
Chromosome numbers and/or ploidy information were c
(a) Assignment of entry numbers and ‘prime’ entries. entered for 90 taxa where previous C-value reports also t/96
TfohrepnruemviboeursinlgistosfothfeDANpApenadmixoutanbtsle(gseeenearbaollvyef)o.lTloawxsatahraet iwnictlhudtheodsethriespionrftoerdmhaetiroen. FaonrdetxhaemCp-lev,alaueCs-vaaglrueeedofc2lo(cid:2)9seplgy /2/229
arranged alphabetically by genus and species with a new for Achillea filipendulina was previously estimated by /29
numberusuallybeingassignedforeachnewspecies. 9
Dabrowska (1992), who also reported that the accession 1
maFdoerf4o2rstpheecsiaemsientshpeecAiepspe(en.dgi.xfmorodreiftfhearennotnceueltsitvimarast,evwarais- sthtuedsiaemdehsapde2cines=in2xth=eA18p.peAnsditxhew1asCv-vearyluseimofila2r(cid:2)8to8tphgatfoorf 62 by
eties,etc.).Inmostcases,theC-valueswereverysimilarand Dabrowska(1992),thechromosomenumberandploidydata gu
e
thedifferentaccessionswereassignedthesameentrynumber s
butweredistinguishedbyletters,sortedherebyincreasing werealsoentered. t on
DNAamounts.Forexample,C-valuesarereportedfortwo (e) Life cycle types. Informationonthetypeoflifecycle 15
cultivarsofCrocuschrysanthus,aspeciesnotincludedpre- forspeciesintheAppendixwaseithertakenfromtheliter- Ap
viouslyinanyofthelistsmentionedabove.Thesmallerof atureorinternet.Thefollowingabbreviationswereused:A= ril 2
thetwovalues,for‘Dorothy’,islistedasentrynumber113a, annual;B=biennial;AP=annual–perennial;P=perennial. 01
A superscript ‘e’ indicates that life cycle information was 9
while the larger, for ‘Blue Pearl’, is 113b. Where several
eitherunknownoruncleartothepresentauthors.
estimates are available for the same species, the ‘a’ value
is automatically chosen in any arithmetical or statistical (f)DNAamountsandconversionfactor.1Cand4Cvalues
calculations.Inthiscontext,asingleestimateforaspecies listedintheAppendixweredeterminedbyappropriatecal-
and‘a’valuesarereferredtoas‘primeentries’. culation of the 2C-value. For 1C-values in megabase pairs
For18speciestheC-valuesbetweenaccessionsdiffered (Mbp) the conversion factor of 1pg = 980 Mb (Cavalier-
by >10 %. Taxonomic heterogeneity, polyploidy and/or Smith,1985;Bennettetal.,2000)wasused.
intraspecific variation are the most likely explanations
(g) Standard species. The abbreviation used to indicate
(especiallywherearangeofchromosomenumbershaspre-
whichofthefivecalibrationstandardswasusedtoestimate
viouslybeenreportedforthespecies),yetuntilchromosome
the 2C-value for each species in the Appendix is given in
counts are made, the cause(s) of the differences remains
Table1.
unknownandthedifferentaccessionsofaspecieswereusu-
allyassignedthesameentrynumber.InsixSeneciospecies (h) Chromosome numbers in Hyacinthus. Brandham
(S.herreianus,S.kleiniiformis,S.mweroensis,S.nyikensis, and West (1993) reported chromosome numbers in
234 Zonneveld et al. — Nuclear DNA Amounts in Angiosperms
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gthistabl Standardgspecies A.americaA.americaA.americaA.americaA.americaA.americaA.americaCliviaA.americaA.americaA.americaA.americaA.americaA.americaCliviaA.americaA.americaA.americaA.americaCliviaA.americaA.americaA.americaA.americaA.americaA.americaA.strictaA.strictaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.americaA.america
n
recedi 4C(pg) .115.323.84.90.59.674.566.336.170.87.154.152.228.884.1160.696.834.932.864.1158.698.654.1298.566.1274.1250.374.354.446.198.202.406.200.224.232.674.552.658.568.46.230.556.616.854.642.778.542
p
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note fount 2C(pg) .58.162.42.45.29.337.283.168.85.43.77.76.114.442.580.348.417.466.432.579.349.327.649.283.637.625.187.177.223.99.101.203.100.112.116.337.276.329.284.23.115.278.308.427.321.389.271
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refer DNA 1C(pg) .29.81.21.23.15.169.142.84.43.22.39.38.57.221.290.174.209.233.216.290.175.164.325.142.319.313.94.89.112.50.51.102.50.56.58.169.138.165.142.12.58.139.154.214.161.195.136 D
ptletters 1C(Mbp) 2,8187,9142,0582,2051,43616,51313,8678,2324,1652,1223,7733,7245,58621,65828,42017,05220,43322,83421,16828,37117,10116,02331,80113,86731,21330,6259,1638,67310,9274,8514,9499,9474,9005,4885,68416,51313,52416,12113,9161,1275,63513,62215,09220,92315,72919,06113,284 ownload
erscri Lifecycleetype PPPPPAPAPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPP ed from
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giosperms Highercgroup EEEEEEEMEEMMMMMMMMMMMMMMMMMMMMMMMMMMMMMEMMEEEEE ic.oup.com
ntentin411an cOrder AsteralesAsteralesAsteralesAsteralesMalvalesRanunculalesRanunculalesPoalesSaxifragalesSaxifragalesAsparagalesAsparagalesAlismatalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAlismatalesAlismatalesAlismatalesAlismatalesAlismatalesAlismatalesAlismatalesAlismatalesAlismatalesAsparagalesAsparagalesAsparagalesAsparagalesRosalesPoalesAlismatalesRanunculalesRanunculalesRanunculalesRanunculalesRanunculales /aob/article-abs
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ndnuclearDNA cFamily AsteraceaeAsteraceaeAsteraceaeAsteraceaeMalvaceaeRanunculaceaeRanunculaceaePoaceaeCrassulaceaeCrassulaceaeAsparagaceaeAsparagaceaeAlismataceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeAraceaeAraceaeAraceaeAraceaeAraceaeAraceaeAraceaeAraceaeAraceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeRosaceaePoaceaeAraceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceae ct/96/2/229/29916
a 2
e, b
.Chromosomenumber,ploidylevel,life-cycletypX bSpecies AchilleafilipendulinaLam.AchilleamillefoliumL.AchilleamontanaSchleich.exDCbAchilleavulgarisAdansoniadigitataL.AdonisannuaL.AdonisvernalisL.AeluropuslittoralisParl.Aeoniumnobile(Praeger)PraegerAeoniumsimsii(Sweet)StearnAgavehartmanniiS.WatsonAgaveportoricensisTrel.exA.BergerAlismalanceolatumWith.AlliumaflatuenseB.Fedtsch.cv.PurpleSensationbAlliumampeloprasumAlliumcepaascaloniumvar.L.AlliumkaratavienseRegelbAlliummolyAlliumnarcissiflorumVill.bAlliumporrumAlliumsativumL.bAlliumschoenoprasumAlliumschubertiiZucc.AlliumsphaerocephalonL.AlliumursinumL.bAlliumvinealeAlocasialongilobaMiq.Alocasialowiigrandisvar.HookAlocasialowiiveitcheivar.HookAlocasiareginulaA.HayAlocasiasanderianaHort.exBullbAlocasiasp.bAlocasiaamazonica·AlocasiazebrinatigrinaK.Koch&Veitchvar.cv.SuperbaAlocasiazebrinareticulataK.Koch&Veitchvar.AloeclavifloraBurch.&ReynoldsAloeecklonisSalm-DyckAloegariepensisPillansAloemicracanthaHaw.AmelanchiercanadensisMedik.AmmophilaarenariaLinkAmorphophalluslewalleiF.Malaisse&P.BampsAnemoneblandaSchott&KotschyAnemoneblandaSchott&KotschyAnemonecoronariaL.AnemonenemorosaL.AnemonepavoninaLam. y guest on 15 April 2019
I
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APP Entrynumb 1b2d3a4a5b6a7a8a9a10a11a12a13b14d15d16l17e18e19a20g21e22b23c24f25e26b27a28a28b29a30a31a32a33a33b34a35a36b37a38a39b40a41c41d42b43c44b
Zonneveld et al. — Nuclear DNA Amounts in Angiosperms 235
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808226628268064892848264826082862823224486642680226264
.9.2.7.8.2.0.9.0.9.1.7.5.1.3.1.9.2.6.5.2.1.4.3.2.5.9.9.7.9.2.2.9.0.8.6.4.2.9.2.3.2.7.4.7.9.8.8.9.9.9.0.3.0.0
785 1117 12327545 11 34 53457343 3 6 11 245333336644
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342 3 1113222 12 26223121 1 3 122111113322
055117965890349526510146589801246211631479298778882821
.0.0.4.2.3.2.4.7.2.2.6.9.5.8.2.2.3.1.1.3.3.1.8.0.1.4.4.1.5.8.8.2.7.2.9.1.5.2.3.3.0.1.6.1.4.9.9.9.9.9.5.5.0.0
221 1 1111 1 13111 1 1 11 1111
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551090161009989597792199401744762771145032593201961519421038891029232388421504202515651689036152399156869058239254038283696862027613504457506555604604847484947898 wn
9,0,4,2,2,2,4,7,2,2,6,8,5,8,2,2,2,1,1,3,2,1,8,0,1,4,4,1,4,7,8,2,7,2,8,1,5,2,3,3, 1,6,1,4,9,9,9,9,9,4,5,9,9, lo
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RanunculalesRanunculalesAsparagalesAlismatalesAlismatalesEricalesAlismatalesAlismatalesAsparagalesPoalesPiperalesPiperalesAsparagalesAsparagalesGarryalesGarryalesPoalesRanunculalesRanunculalesAlismatalesAsparagalesAsteralesAsparagalesPoalesLamialesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAsparagalesAlismatalesPoalesAsparagalesAlismatalesRanunculalesAsparagalesEricalesEricalesGentianalesGentianalesAsparagalesAsparagalesAsparagalesRanunculalesRanunculalesRanunculalesRanunculalesRanunculalesRanunculalesRanunculalesRanunculalesRanunculales ob/article-abs
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RanunculaceaeRanunculaceaeAsparagaceaeAraceaeAraceaeMyrsinaceaeAraceaeAraceaeAsparagaceaePoaceaeAristolochiaceaeAristolochiaceaeXanthorrhoeaceaeAsparagaceaeGarryaceaeGarryaceaePoaceaeBerberidaceaeBerberidaceaeAraceaeOrchidaceaeCampanulaceaeAsparagaceaePoaceaeScrophulariaceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeXanthorrhoeaceaeButomaceaePoaceaeAsparagaceaeAraceaeRanunculaceaeAsparagaceaeTheaceaeTheaceaeApocynaceaeApocynaceaeAlliaceaeAsparagaceaeAsparagaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceaeRanunculaceae ct/96/2/229/2991
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bAnemoneranunculoidesbAnemonetrifoliaAnthericumliliagoL.Anubiasbarterinanavar.SchottAnubiasbarteriSchottbArdisiacrenataArisaemaciliatumH.LiArisarumvulgareTarg.Toz.bArthropodiumcirrhatumbArundodonaxAsarumeuropaeumL.Asarumsplendens(Maekawa)C.Y.Cheng&C.S.AsphodelineluteaReichb.bAspidistraelatior(variegated)AucubajaponicaThunb.AucubajaponicaThunb.cv.RozannieAvenasativaL.BerberisthunbergiiDC.cv.AtropurpureaNanabBerberisvulgarisbBiarumtenuifoliumBletillastriataRchb.f.BrighamiainsignisA.GraybBrimeuraamethystinabBromusmollisBuddlejadavidiiFranch.BulbineasphodeloidesSpreng.Bulbinebulbosa(R.Br.)Haw.BulbinefallaxPoelln.BulbinefrutescensWilld.BulbinehookeriKunthBulbinemargarethaeL.I.HallBulbinellalatifoliaKunthButomusumbellatusL.bCalamagrostisepigeiosCalibanushookeriiTrel.exA.BergerCallapalustrisL.CalthapalustrisL.CamassiacusickiiS.WatsonCamelliasinensisassamicaKuntzevar.CamelliasinensisKuntzeCatharanthusroseus(L.)G.DonCeropegiawoodiiSchltr.ChlidanthusfragransHerb.bChlorophytumcomosumChlorophytumhoffmanniiEngl.ClematisarmandiiFranch.cv.BowlofBeautyClematisarmandiiFranch.cv.EnhamStarClematisarmandiiFranch.cv.AppleBlossomClematisarmandiiFranch.cv.LittleWhiteCharClematisarmandiiFranch.cv.SnowdriftClematisflammulaL.ClematisforsteriJ.F.Gmel.bClematismontanabClematisvitalba 15 April 2019
5a6a7a8a8b9a0a1a2a3a4b5a6a7b8a8b9f0c1b2a3a4a5a6a7a8a9b0b1a2a3a4a5c6a7a8a9c0b1b1c2b3b4a5a6a7a7b7c7d7e8a9a0a1b
444444555555555556666666666777777777788888888888888899
236 Zonneveld et al. — Nuclear DNA Amounts in Angiosperms
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Standardgspecies A.americA.americA.americA.strictaA.americA.americA.americA.strictaA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americA.americCliviaA.americA.americA.americA.strictaA.americA.americA.americA.americA.americ
4C(pg) .780.112.134.308.136.150.374.42.1010.1032.804.830.838.40.38.56.37.40.36.45.46.36.262.156.162.442.36.208.206.590.554.176.726.270.134.156.248.150.62.850.530.356.478.328.1470.1428.1346
fount 2C(pg) .390.56.67.154.68.75.187.21.505.516.402.415.419.20.19.28.19.20.18.22.23.18.131.78.81.221.18.104.103.295.277.88.363.135.67.78.124.75.31.425.265.178.239.164.735.714.673
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DNA 1C(pg) .195.28.34.77.34.38.94.11.253.258.201.208.210.10.10.14.09.10.09.11.12.09.66.39.41.111.09.52.52.148.139.44.182.68.34.39.62.38.16.213.133.89.120.82.368.357.337
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Cbp) 110744283546332675163029745284698335531985936372907975877098127872419822969829872096047455573312787615283812076675529825985722711036015986977 wn
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A.PPENDIX bSpecies CliviaminiataRegelCocosnuciferaL.ColchicumautumnaleL.CommelinadianthifoliaDelileCornusmasL.Cortaderiaargentea(Nees)StapfCotinuscoggygriaScop.CraterostigmaplantagineumHochst.Crinumasiaticumjaponicumvar.BakerCrinumasiaticumsinicumvar.BakerCrinumbulbispermumMilne-Redh.&SchweickerdtCrinummooreiHook.f.Crinumnatanscalamistratumvar.BakerCrocosmiaaurea(Hook.)Planch.Crocosmiafucata(Herb.)M.P.deVosCrocosmiafucata(Herb.)M.P.deVos(pleatedform)Crocosmiamasonorum(L.Bolus)N.E.Br.Crocosmiapaniculata(Klatt)GoldblattCrocosmiapottsii(Baker)N.E.Br.bCrocosmiacrocosmiiflora·bCrocosmialatifolia·bCrocosmiapaniculata·Crocusangustifoliusflavus(largeyellow)·CrocuschrysanthusHerb.cv.DorothyCrocuschrysanthusHerb.cv.BluePearlbCrocusvernus(whiteform)CryptocorynecrispatulaEngl.CyanotissomaliensisC.B.ClarkebCyclamenhederifoliumCyrtanthuselatus(Jacq.)TraubCyrtanthusmontanusR.A.Dyercv.FirebirdDactylisglomerataL.Dichelostemmaida-maiaGreeneDioscoreaelephantipesEngl.bDoryanthespalmeriDracunculuscanariensisKunthDracunculusvulgarisSchottDrimiopsismaculataLindl.&Paxt.Eichhorniacrassipes(Mart.)SolmsbElymusarenariusElymusjunceusElytrigiajunceaFisch.()=EranthiscilicicaSchott&KotschyEranthishyemalisSalisb.EranthispinnatifidaMaxim.Erythroniumhybridcv.NanaErythroniumhybridcv.PagodabErythroniumcalifornicumcv.WhiteBeauty st on 15 April 2019
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ntryumb 2c3c4a5b6a7a8a9a00a00b01a02a03a04a05a05b06a07a08a09a10a11a12a13a13b14b15a16a17b18a19a20g21a22a23b24a25b26a27b28a29a30a31b32a33a34a35a
En 99999999111111111111111111111111111111111111111
Zonneveld et al. — Nuclear DNA Amounts in Angiosperms 237
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LiliaceaeLiliaceaeEscalloniaceaeAlliaceaePoaceaeAsparagaceaeCrassulaceaeCrassulaceaeGunneraceaeGesneriaceaeAraliaceaeAraliaceaeAraliaceaeAraliaceaeAraliaceaeAraliaceaeAraliaceaeAraliaceaeMelanthiaceaeIridaceaeAsparagaceaeAlliaceaeAlliaceaeAlliaceaeAlliaceaeApocynaceaePoaceaePoaceaeArecaceaeApocynaceaeCannabaceaeCannabaceaeCannabaceaeAsparagaceaeAsparagaceaeAsparagaceaeAsparagaceaeAsparagaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceae HydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceaeHydrangeaceae t/96/2/229/29916
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bErythroniumdens-canisbErythroniumjaponicumEscallonialangleyensisVilm.&BoisEucharisgrandifloraPlanch.&LindenFestucaovinaL.cv.GlaucaFurcraeaselloaK.Koch(yellowedge)´GraptopetalumbellumMoran&J.MeyranGraptopetalummacdougalliiAlexander´GunneramanicataLindenexAndreHaberlearhodopensisFriv.bHederaalgeriensiscv.GloiredeMarengobHederacanariensisHederacaucasigenaPojark.cv.TanjaHederacolchicaK.Kochcv.SulphurHeartHederacolchicaK.Koch(yellowedge)HederahelixLcv.Erecta.HederahelixLcv.GoldHertz.HederahibernicaHort.bHeloniopsisorientalisHermodactylustuberosaMill.Hesperaloeparviflora(Torr.)J.M.Coult.HippeastrumcybisterBenth.&Hook.f.HippeastrumelwesiiC.H.WrightHippeastrumpapilio(Ravenna)J.VanScheepencv.ButterflyHippeastrumreticulatumstriatifoliumvar.Herb.Hoodiagordonii(Masson)SweetexDecne.HordeummurinumL.HordeumvulgareL.cv.SultanHoweaforsterianaBecc.Hoyacarnosa(L.)R.Br.HumulusjaponicaSiebold&Zucc.bcordifoliaHumuluslupulusvar.bHumuluslupuluscv.BramblingCrossHyacinthusorientalisL.cv.WhitePearlHyacinthusorientalisL.cv.PinkPearlHyacinthusorientalisL.cv.AnnaMarieHyacinthusorientalisL.cv.DelftBlueHyacinthusorientalisL.cv.L’InnocenceHydrangeaanomalapetiolarisssp.D.Doncv.CordifoliaHydrangeaanomalapetiolarisssp.D.DonHydrangeaarborescensL.Hydrangeaasperasargentianassp.Buch.-Ham.exD.DonHydrangeaasperarobustassp.Buch.-Ham.exD.DonHydrangeaasperastrigosassp.Buch.-Ham.exD.DonHydrangeaasperaBuch.-Ham.exD.Doncv.VillosaHydrangeaaspera?Buch.-Ham.exD.Don(fromTaiwan)HydrangeaheteromallaD.DonHydrangeaheteromallaBuch.-Ham.exD.Doncv.NepalBeautyHydrangeahirtaSieboldHydrangeaintegrifoliaHayataHydrangeainvolucrataSieboldbHydrangeamacrophylla(diploid)bHydrangeamacrophylla(triploid) y guest on 15 April 2019
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238 Zonneveld et al. — Nuclear DNA Amounts in Angiosperms
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4C(pg) .140.43.80.83.95.86.88.386.646.498.168.224.528.282.410.160 .432.148.152.110.122.226.224.234.102.178.190.548.1916.1762.1546.1690.1864.106.422.38.114.74.978.86.198.612.694.668.822.678
fount 2C(pg) .70.22.40.42.48.43.44.193.323.249.84.112.264.141.205.80 .216.74.76.55.61.113.112.117.51.89.95.274.958.881.773.845.932.53.211.19.57.37.489.43.99.306.347.334.411.339
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(CoA.PPENDIX bSpecies bHydrangeapaniculataHydrangeaquercifoliaBartram(Tennesseeclone)bHydrangeascandensliukinensisvar.bHydrangeascandensHydrangeascandens?cv.AngustataHydrangeaserrataSer.Hydrocharismorsus-ranaeL.Ipheionuniflorum(Lindl.)Raf.IrisdanfordiaeBoiss.bIrisgermanicaIrislacustrisNutt.IrispseudacorusL.bIrispumilaIrisreticulataM.Bieb.IrisreticulataM.Bieb.cv.HarmonyIrissintenisiiJankaKniphofiahybrida(OttoMannexGumbletoncv.LittleMaidLachenaliahybridcv.NamaquaLachenaliahybridcv.RomaudLactucasativaL.cv.IJsbergbLaurusnobilisLavandulaofficinalisChaixLedebouriacooperiadlamiivar.(Hook.f.)JessopLedebouriasocialis(Baker)Jessopcv.ViolaceaLewisiacotyledonRobinsonLewisianevadensis(A.Gray)B.L.Rob.Lewisiapygmaea(A.Gray)B.L.Rob.LiliumamabilePalib.bLiliumcanadensebLiliumcandidumLiliumeuxanthum(W.W.Sm.&W.E.Evans)SealyLiliumformosanumWallacebLiliummartagonLimoniumvulgareMill.LiriopemuscariL.H.Bailey.LoniceranitidaE.H.WilsonbLonicerapericlymenumLunariabiennisMoenchLycorisaureaHerb.Magnoliakobusstellatavar.Maxim.MagnolialiliifloraDesr.Maianthemumbifolium[DC.]MaianthemumcanadenseDesf.Maianthemumdilatatum(Wood)A.Nelson&J.F.Macbr.Maianthemumjaponicum(A.Gray)J.V.LaFrankieMaianthemumracemosumLink guest on 15 April 2019
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ntryumb 77b78b79a79b79c80a81a82a83a84a85a86b87a88a88b89a90a 91a91b92d93a94a95a96a97a98a99a00a01a02a03a04b05a06a07b08a09b10a11a12b13a14a15a16a17a18a
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