Copyright (c) American Society for Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 uosoue n h rsneo ogcanpolyunsatu- chain long of presence the and pen- ovules monophyletic central free dulous supported include order strongly the for a Synapomorphies clade. is analyses phylogenetic uncertainty that previous Despite show placement, global previ- (1984a). its as Sleumer about older, by are most suggested they in portrayed ously that as but rosids, classifications, derived more traditional to Santalales that related suggests not position asterids are This 2003). and 2000, al. in- rosids, et also (Soltis Gunnerales, that polytomy Caryophyllales, core a the from cludes among emerges order it this however, placed , clearly has phy- work Molecular logenetic Viscaceae. Olaca- and Santalaceae, Misodendraceae, Opiliaceae, Loranthaceae, clas- ceae, typically families: are six that in species 2200 in- sified and It genera members. 160 parasitic ca. and cludes woody both with angiosperms iul hw Ncrn n uf19;Ncrn ta.1998), al. et pre- Nickrent As 1996; Duff “Olacaceae”. and of (Nickrent small- clades shown chloroplast eight viously nuclear of and from existence (=18S) the data mented rDNA sequence in (SSU) used genera Malécot 28 subunit and and the of Nickrent family 17 1998; sampled the which al. study, latter et The Nickrent 2001). (Nickrent 1996; polyphyletic Duff is and Olacaceae analyses that phylogenetic evidence molecular 1955; provided recently, Reed More into 1948; 1968). split Fagerlind be Kuijt 1935; could (Sleumer that families assemblage multiple ex- polyphyletic a this Ola- consider as Historically, to caceae workers 1984). many 1982; prompted Oever has Baas variation den treme 1980; van Lobreau-Callen mor- 1984b; example and Sleumer for palynology (see anatomy, in phology heterogeneity In Olacaceae show of 1969). genera also Kuijt all modes, 1948; nutritional Fagerlind diverse Engler to 1935; addition both 1894; Sleumer (Engler 1924; contains Gilg species it and nonparasitic because and “primitive” parasitic most root the as regarded acquisi- habits. evolutionary parasitic the various study of to group tion sandal- ideal the an reason, autotrophs, are this includes For woods parasites. it stem because and parasites angiosperm root of seen order modes nutritional any of angiosperms. in range other broadest in fea- the absent both have or acid), Santalales rare santalbic otherwise are or that ximeninic tures (e.g. acids fatty rated oyih 08b h mrcnSceyo ln Taxonomists Plant of Society American the by 2008 Copyright © Botany Systematic mn h atlla aiis lcca a ogbeen long has Olacaceae families, santalalean the Among of group largest the is (Santalales) order sandalwood The asmn n aeinaaye aersle ee lds well-suppor clades, seven resolved subs chloroplast have in and analyses evolved rDNA Bayesian that SSU and innovations (nuclear parsimony many data sequence for DNA ground obtained staging This the perianths. monochlamydous represents example and it For dichlamydous with classification. members their and regarding trees, confusion much caused cally iimapast aefrteovdi h ld containing clade the in evolved autotrophic. first have to appears sitism 1 Keywords— Abstract— M42SGH ntttNtoa ’otclue NA nvAgr,2ru 2 Univ-Angers, INRA, d’Horticulture, National Institut SAGAH, UMR462 a oecoeyrltdt ioedaeeand Misodendraceae to related closely more was oeua hlgntcRltosiso lcca n eae Santalal Related and Olacaceae of Relationships Phylogenetic Molecular 20) 31:p.97–106 pp. 33(1): (2008), 2 eateto ln ilg,Suhr liosUiest,Carbondale, University, Illinois Southern Biology, Plant of Department staiinlycrusrbd h aiyOaaeecnan morphol a contains Olacaceae family the circumscribed, traditionally As hools N,Oaaee aaiim hlgn,rbsmlDA Sant DNA, ribosomal phylogeny, parasitism, Olacaceae, DNA, chloroplast 3 uhrfrcrepnec ([email protected]) correspondence for Author aéyMalécot Valéry omnctn dtr nraSchwarzbach Andrea Editor: Communicating rbcL docu- , 1,3 n ailL Nickrent L. Daniel and rbcL hra ldsbtenta n n h ugopapa ob entirely be to appear outgroup the and one that between clades whereas 97 and lcca oti aaie n oprsts lmiglaa and lianas climbing nonparasites, and parasites contain Olacaceae , n aéo 01 aeilietfe as identified material 2001) Malécot genus and The 2002). Malécot by in represented done was as genus, distinct DAwsapiidadsqecduigte1 owr (5 forward SSU 12 (1994). the Nickrent using sequenced in and described amplified as was tissue rDNA leaf preserved herbarium or Saxi- out- Cornales Caryophyllales, Solanales. Vitaceae, 15 Proteales, Brassicales, the of eudicotyledons, Rosales, of members Trochodendraceae, the fragales, Because from within 1). selected (Appendix were Santalales groups sampled of were position Santalales ambiguous of taxa 43 total, be to proved that the is and exception defined (the well paraphyletic be are genus to genera type proposed the or been ever that of two have in availability few include justified very by also to partially is genera determined but spe- was larger material single sampling for a This made by species. were represented more . were attempts among species whereas placed eight best cies, than is it fewer that with showed Genera (2002) Malécot by study cal unsuccessful. proved oese l 06.TermiigSnaae r ersne ysxgen- Loranthaceae, six of by represented four are Opiliaceae, Santalales of remaining The era 2006). al. et Rogers of specimens extrac mono- herbarium to rare Attempts available 1976. extremely in from this once, and only DNA recollected 1984 been in has genus described specific was latter Octokne- except The into and cluded. genera segregated Erythropalaceae, Olacaceae been , All times as maceae. at such have families that distinct those including sampled, were htalesnilyeegdfo oyoya h aeof base the at polytomy a from emerged such essentially poor all was clades that seven remaining the among Resolution o aiu ldso Olacaceae. synapomorphies of morphological clades various identify for 3) within groups and various basal of Olacaceae, all monophyly among the assess relationships 2) these phylogenetic Santalales, use clarify to 1) is goal to Our data Santalales. compre- basal nearly of a sampling among hensive relationships assess to genes roplast 2004). as al. et well (Malécot (as Olacaceae resulted Olacaceae paraphyletic families) a of santalalean in genera other the 28 from all char- representatives morphological sampled 80 and used that acters study cladistic A order. the nisonfmly copica lm,ta o loin- also now that Blume, familly, cludes own that its consider in we follow- (2002), and al. work et Judd this ing From Olacaceae. to than Loranthaceae C n Sequencing— and PCR aooi Sampling— Taxonomic nti td,w sdsqecsfo ula n chlo- and nuclear from sequences used we study, this In e ymlclradmrhlgclcaatr.Ro hemipara- Root characters. morphological and molecular by ted matK qetgop.Tepeetmlclrpyoeei td has study phylogenetic molecular present The groups. equent aiyi aams ntesnawo re Snaae) thus (Santalales), order sandalwood the in basalmost is family Arjona o l u w fte2 eeai hsgop Maximum group. this in genera 28 the of two but all for ) gclydvreasmlg fgnr hthshistori- has that genera of assemblage diverse ogically lxemirnensis Olax ncls papuana alales. hc sprpyei when paraphyletic is which and Brachynema M eNte -94 nes France Angers, F-49045 Nôtre, Le e Quinchamalium TRASAND ATERIALS 2 N a xrce rmslc e dehydrated gel silica from extracted was DNA wnysxgnr n 2seiso Olacaceae of species 32 and genera Twenty-six lios69160 U.S.A. 62901-6509 Illinois Mlct20;MlctadDbisn2004; Dubuisson and Malécot 2002; (Malécot a xlddbcuetemorphologi- the because excluded was hra norfrtsuy(Nickrent study first our in whereas , M .casearioides A. Misodendrum DradNcrn 2008). Nickrent and (Der ETHODS Schoepfia Dulacia Harmandia and Douradoa srcgie sa as recognized is and , ( caze l 3620 al. et Schatz es Anacolosa sbs placed best is and Schoepfia Ј TCTGC -TCC Douradoa eein- were shere is and , .In ) t Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 in fti oe eg T + GTR (e.g. model this GT analyzed of in were sions resulted positions all third two vs. but the For second separately, partitions. and gene first (GT three genes, the model of protein-coding each runs Reversible for so MrModeltest Time by 0.001, selected probability below General posterior was the The in cases likelihood trees distribution. of all −ln number in the the increasing runs in combined were between stationarity variance by to set The one determined was score. was than priors the burn-in partitions), more of The rate codon When the vary. and and analysis. unlinked partitions were the gene estimations parameter all (separate of was analyzed distribution to part was prior assigned as Dirichlet partition estimated were a which and probabilitites as for assigned prior frequencies estimated uniform state were the analysis; generations, parameters million except the Model 100 five run. of every for each part saved performed trees were were 50,000 parameters each producing indepen- and two chains BI, Trees four For were generations. min./replicate. with partition 1 rDNA to analyses se- limit SSU addition dent time the a simple for setting with by estimates replicates obtained Bootstrap 100 TBR. from and obtained quence (Felse values were support 1985) (BS) the stein Bootstrap using indices. estimated retention was and recon- Homoplasy consistency bisection algorithm. Tree by swapping the selected (TBR) and nection searches, replicates), model heuristic (200 used sequence a analyses addition MP with random 2004). using al. 2003) et performed Huelsenbeck (Nylander was MrModeltest BI and whereas (Ronquist were 2002) analyses MrBayes (Swofford MP missing. PAUP* as in coded combined were and performed gaps individual analyses, for all performed In were partitions. analyses (BI) inference deposited ian were taxa Santalales 1). GenBank for (Appendix from sequences obtained GenBank were generated with taxa newly outgroup 15 the the Sequences and for S1843). genes number 3-gene three (study all The TreeBASE for matrix. from the available of is 12.9% alignment constituted indels and sequence missing l neswr rae smsigdt.SUrN from rDNA SSU data. missing as frame. treated sequence were maintaining while indels introduced All were indels and quences the lcdi eA Rmat20)adaindmnal.Fraligning then For manually. were aligned sequences and 2004) and Edited (Rambaut 2006). Se-Al in or Groothius placed 2000), and Codes (Gene (Griekspoor 4.1 4Peaks Sequencher 1997), (Parker Navigator Biosys- Sequence Applied sequencer, DNA automated manufacturer to automated using according 377 conducted tems) was Prism Adel- sequencing (ABI Inc. Direct Separations, methods (Princeton Jersey). Sephadex New with phia, columns Centri using spin purified 100 were manufacturer products Sep reaction the sequencing to cycle Pro- according The tocol. Easy, Wisconsin) pGEM-T Madison, (Promega For Corp., performed Georgia). mega was Liburn, commer- cloning Inc., a TA BioTek using samples, Omega purified gels, some Kit, and agarose Cycle-Pure excised, (EZNA 1.0% size kit on appropriate cial purified the were of products bands PCR the DNA. genomic of ng 50 aio,Wsosn 0MTi C,5m C,p .) . MMgCl mM 1.5 TAT 8.3), pH CTT KCl, ATA 50mM HCl, TAT Tris AAG 10mM 1 Wisconsin; (TCG included: Madison, reactions rev specifi- amplification 1316 PCR one Typical study, and TCG). used, this were for (1997) designed Liang cally and Hilu by developed primers 6 9 n 0sqecsfrSUrDNA, SSU for sequences 40 and 39, 26, -ee1 ,9 .87036 .87460279711,180 171 711 202 2,799 4,690 1,426 0.5897 1,799 0.3968 0.4837 0.6209 0.3796 5,691 0.538 10 842 3-gene 1,061 matK rbcL rDNA SSU 33 [Volume BOTANY SYSTEMATIC (5 (5 reverse forward the 3 with T- TGT GC-3 CCT TAT TCA TAS CAG 98 hlgntcReconstruction Phylogenetic euneAlignment Sequence T oa f15nwsqecswr bandi hsstudy: this in obtained were sequences new 105 of total A ␮ ABLE matK dNTP M matK and .Gn iest ttsisfrtevrospriin n obnddata combined and partitions various the for statistics diversity Gene 1. h uloiesqecswr rnltdit mn cdse- acid amino into translated were sequences nucleotide the , Ј aastlacked set data TGTAAGATGGCGAG-3 CCG GGG ATT AAG TAA -TAG matK ’ ,1ui a oyeae 0.4 polymerase, Taq unit 1 s, Ј rmr.The primers. ) o re reLnt ..CI-RI o hr.Cnev hr.Vral U Variabl. Chars. Conserv. Chars. No. R.I. C.I.- C.I. Length Tree Trees No. from 0 ,2 .95045069 ,3 0 7 653 276 506 1,435 0.6199 0.435 0.4995 3,023 101 Ј – — ,3 .62039 .731408420346 230 864 1,440 0.5733 0.3691 0.4682 1,636 9 T C C A C A AC-3 GAR ACA CAA CCA TCA ATG Engomegoma lcrpeormflswr dtduigeither using edited were files Electropherogram Engomegoma Ј n 79rvre(5 reverse 1769 and ) — rbcL R ⌫ ’ aiu asmn M)adBayes- and (MP) parsimony Maximum R+I+ protocols. s ESULTS orSYM+I+ eentotie.Tkntogether, Taken obtained. not were eewsapiidadsequenced and amplified was gene . ossec ne,CI-=cnitnyidxmnsuifraiec uninformative minus index consistency = C.I.- index, consistency = C.I . ⌫ ␮ rsihl escmlxver- complex less slightly or fec rmradc.30 ca. and primer each of M ⌫ Ј Ј rmr.For primers. ) CCCACGAAA CGG CTA -CAC ). rbcL × ufr(Promega, buffer R+I+ and Cathedra matK Ј matK n 3 and ) ⌫ ’ pro- s was ) re- , the , rbcL and n- 2 Ј - , s o S DAadlws in lowest and rDNA SSU the high- for the est expected, to 9.4%) be (171, characters rDNA in would SSU est informative in As lowest parsimony the 1. from and ranged Table variable in of 3-gene shown number concatenated is the set and data partitions gene separate three related more or two among taxa. shared were is, others length. that of in number synapomorphic; codons a whereas three autapomorphic to were one Some from ranging generally tions, aeyeuvln,btlwrthan lower and but rDNA The equivalent, sites). SSU mately informative for of indices number consistency higher its given expected Indel aligned. excluded unambiguously were be rDNA, in SSU not events the could of they region V4 because the in all tions, o h sie ftete uhta h nru a not was ingroup of support the BS that received clades such 45 tree of five the Only obtained of monophyletic. was “spine” resolution little the rDNA, SSU for For 1. Fig. in pared SSU from were positions 1,815 from these, and 1,440 ingroup rDNA, Of 43 taxa. for positions outgroup aligned 15 4690 had set data rDNA, gene) SSU concatenated The spectively. n Po . Fg ) oeae(S=7)t ih(P=1.0) = clade. (PP asterid high the to between and 71) relationship = Santalales sister-group 2) (BS a for Moderate (Fig. obtained 3). was support (Fig. support mono- 1.0 BS of a 100% PP resolved with a BI and and A) MP dis- (clade Both be Santalales 3). will phyletic 2, two Figs. the on relevance thus particular A-J of (labeled tree, clades MP upon focusing the together, as cussed many clades recovered 3) same (Fig. BS of the tree received BI 30 of The consensus clades, greater. total or 52 strict 90% the of most Of the support 2. ten Fig. steps, in yielded shown 5,691 is matrix which to- of 3-gene analyzed trees the of and parsimonious reso- analysis concatenated of MP were degree gether. the partitions in the only lution, differed generally and topology within clades supported. of highly number a support were and BS Olacaceae 99% greater. was or the clade ingroup 90% of the of 24 for support with BS partitions receiving three clades the 46 among resolution parsi- of of level number the higher sites, its informative With the mony of resolved. spine poorly the BS all along were clades 50% tree trees the < rDNA, shortest SSU received with clade the As this support. Although ingroup, monophyletic greater. a or show 90% of support the For genera. support the obtain Opiliaceae, to as exists such clades, signal several sufficient for but greater, or 90% ld,adtecaeta includes that clade the and clade, eedvriysaitc olwn Paayi o the for analysis MP following statistics diversity Gene h oooiso h niiulM eetesaecom- are trees gene MP individual the of topologies The staiinlydfnd(..including (i.e. defined traditionally As in conflicting not were trees gene individual the Because es h S DAdt e akdsqecsof sequences lacked set data rDNA SSU The sets. matK matK 63 55) h ubro hrettesi high- is trees shortest of number The 45.5%). (653, rbcL eeotnrcgial stne duplica- tandem as recognizable often were rbcL, atto,1 fte4 ldsrcie BS received clades 46 the of 19 partition, n ,3 from 1,435 and matK aatr,RI eeto index. retention = R.I. haracters, rbcL eepoie h highest the provided gene matK Anacolosa (not ifr.Vral Inform. Variabl. ninform. . matK Olax rbcL rbcL matK Schoepfia Cathedra hrenposi- Thirteen . n orother four and / eeapproxi- were and smgtbe might as ,Olaca- ), and matK Coula (3- ; Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 l lcca,Ot=outgroups. = Opi Out are: Olacaceae, abbreviations Taxon = branches. Ola the above given are replications) 08 MAL 2008] F IG .Mxmmprioytesfrtetreidvda eepriin,SSU partitions, gene individual three the for trees parsimony Maximum 1. . É O IKET LCCA HLGN 99 PHYLOGENY OLACACEAE NICKRENT: & COT plaee o oataee i ioedaee c Schoepfiace = Sch Misodendraceae, = Mis Loranthaceae, = Lor Opiliaceae, = rDNA, rbcL and , matK otta ausgetrta 0 (100 50% than greater values Bootstrap . ae, Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 0 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 100 icsini et.Txnabeitosaea nFg 1. Fig. in giv as are are replications) abbreviations (100 Taxon text). 50% in than discussion greater values Bootstrap Santalales. F IG .Src osnu ftnms asmnostesotie rmte3-ge the from obtained trees parsimonious most ten of consensus Strict 2. . naoetebace.Crldltesietf ldso lcca .lat s. Olacaceae of clades identify letters Circled branches. the above en edt e cnaeae ula S rDNA, SSU nuclear (concatenated set data ne rbcL and , matK o basal for ) (see . Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 dniycae fOaaees a.(e icsini et.Txnabbre Taxon text). in discussion (see lat. s. Olacaceae of clades identify ldsaogtesieo h Pte.CaeB(S=100, = with (BS genera B arborescent Clade six of tree. of number composed a MP for is the support 1.0) of BS = spine low PP poly- by the the and along by tree clades evidenced among BI as the relationships resolved, for in high, tomy always support not generally Although were is 3). them 2, clades (Figs. seven the lineages these of seven core of the posed and without analyses (i.e. these family in polyphyletic are ceae MAL 2008] F IG .Bysa reotie rmte3gn aastfrbslSnaae.P Santalales. basal for set data 3-gene the from obtained tree Bayesian 3. . Schoepfia saprpyei ru com- group paraphyletic a is ) É O IKET LCCA HLGN 101 PHYLOGENY OLACACEAE NICKRENT: & COT Scoro- itosaea nFg .Saea otmgvstenme fsubstitutions of number the gives bottom at Scale 1. Fig. in as are viations ld B 6 P=10 opie orseisof species genera four monospecific comprises two 1.0) the = and PP 66, = (BS BI, C Clade With MP. clade, ing next Engomegoma The stylidium, resolved. con- these not of of were first species The three clades. tains well-supported two of posed borneensis docarpus seirpoaiiisaegvnt h ih fec oe ice letter Circled node. each of right the to given are probabilities osterior eovda itrt ld ute com- further clade a to sister as resolved Strombosiopsis and Diogoa Maburea ssse to sister is eeas neovdus- unresolved also were hs interrelationships whose tobsoss Tetra- Strombosiopsis, and Tetrastylidium e site. per s . . Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 srsle ihmdrt upr B 7 P=05)as 0.53) = PP 77, = (BS support to moderate sister with genus resolved root-parasitic pantropical is This Olacaceae. of member a I Clade of polytomy. composed a 1.0), = from PP arising 100, as = viewed (BS be should to of J only none and support, sister Because strong as probability. received it posterior nodes shows low these BI a with and but 50% J than from clade less BS separate with as J and classified placed been analysis MP frequently Olacaceae. has that taxon Heisterieae. tribe ( in placed latter been the and Anacoloseae tribe as ognized plaeealocri h etr hemisphere. western the in occur all followed Opiliaceae basalmost, is Asia, well- by lianas, South-East a from of shrub 1.0), Opiliaceae, genera notypic Within = 10 trees. small PP of and shrubs, family 100, pantropical = J characterized (BS clade within Opiliaceae clade major represents second The (mistletoes). parasites The America. South BI. or MP by supported not is genera Olax nochiton, eua re,sc si iketadMal and Nickrent mo- in published as previously re- such with trees trees, congruent The lecular are sampling. here taxon ported comprehensive with caceae otenArc and parasite. stem Africa or root southern a either be just parasite can that root by toe The represented by family. followed the here basalmost in 1.0), is genera = 73 the PP of 99, four = (BS Loranthaceae thedra lianescent the for obtained also is support Relatively 0.95). PP Erythropalum = and PP 66, BS = (BS low support moderate to low with monophyletic A 0 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 102 aeicesdrslto.Ti mrvmn,stemming improvement, This resolution. increased have w elspotdsblds h is with first the contains subclades, clade well-supported The accrescent two Discussion). unusual (see show fruiting which upon of structures many trees, large to small the 1.0), to = sister PP is 100, clade = this (BS and F clade For clas- pantropical Couleae. been traditionally tribe have in that sified = taxa (BS arborescent E of group Clade contains a 2). 1.0) (Fig. = MP PP with 100, supported) not (but sister as clas- taxa Opiliaceae. of and Schoepfiaceae, Misodendraceae, remainder Loranthaceae, as the well BI as contains Olacaceae the clade in sified on This (1.0) 3). probability (Fig. posterior tree high with resolved clade. was C plus (0.69) B PP monophyletic BI low a a for and obtained (52%) individual support is the BS of MP any weak on trees, seen gene not is topology the Although 2). ld scmoe oeyof solely composed is H Clade mn h ld aa only taxa, J clade the Among hssuyrpeet h is oeua hlgn fOla- of phylogeny molecular first the represents study This ld B 0,P .)i opsdo i eeaof genera six of composed is 1.0) = PP 100, = (BS G Clade occur do but 3) (Fig. BI with unresolved are F and E Clades but 66%) (i.e. support BS MP poor only received D Clade Agonandra swl upre,bttemnpyyo h atrtwo latter the of monophyly the but supported, well is and , stieAptandreae. tribe as ) Misodendrum and Phanerodiscus Ximenia ssse othe to sister as Ongokea rmtenorpc.Termiigsampled remaining The neotropics. the from Heisteria ssse to sister is Schoepfia/Misodendrum h omrgophsotnbe rec- been often has group former The . nara aaiefo aaoi in Patagonia from parasite aerial an , Tupeia n h eodwith second the and D Gaiadendron sspotda itrto sister as supported is ISCUSSION Maburea rmNwZaadaeaerial are Zealand New from Curupira Schoepfia reedmct China. to endemic tree a , Ptychopetalum Octoknema plus , nabrsetmistle- arborescent an , Octoknema Coula, rmSuhAmerica South from a enconsidered been has Heisteria ssse ocae I clades to sister as Lepionurus ld ssse to sister is clade a traditionally has Moquiniella é and pada Chau- , o 20) but (2001), cot nenigmatic an , Anacolosa Aptandra , Dulacia, n ldsI clades and ld (Fig. clade Maburea Nuytsia mo- a , from , and and Ca- , and oeoa Strombosiopsis gomegoma, thedra ( Olaceae six Couleae, recovered Ximenieae, analyses (tribes): both Lo- clades to Olacaceae, clade this Within of ranthaceae. relationship sister and the Misodendraceae and between Schoepfiaceae, to relationship failed sister tree morphological the The capture de- trees. Olacaceae more both autotrophic a on position Opiliaceae the rived and with position basalmost example fami- be the for the occupying can among similar, dissimilarities relationships is of and lies pattern similarities overall The some Mal noted. in (2004), reported al. cladogram the et with molecu- compared the is When tree Santalales. lar basal major the more in of lineages of understanding phylogenetic addition our the increased has and data, sampling sequence taxon greater both from etrshv e aywrest isolate these to Indeed workers tendrils). many led axillary have with features of climber affinities woody (a the habit characters, such on it Erythropalum Based link Couleae. 1982) (Baas tribe anatomy to leaf and 1982) 1980, breau-Callen teria, other the for topologies genera. conflicting two gave analyses different relation- sister a support between data ship morphological and 3) apiculate Fig. of tree, synapomorphy (Mal the connectives have anther taxa Mal These in BS data. (85% 2004) morphological and BS) (94% lecular genera, rastylidium paucispecific four of clade l ee lccoscae;ti utb ett uuework. future to left be of must this presence among clades; olacaceous the relationships seven discussing clades, all fully advanced precludes more tree from polytomies progression to BI general and basalmost the 2) of the indication (Fig. some Al- tree give clades. MP 3) (Fig. parsimonious the rel- most integrate characterize the will that we though data section following morphological the morphological evant the In on low. nodes was these tree for analy- support two however, the between ses; differed tribes these among tionships eeai hscae ase l 19)sae htwood that stated (1992) al. et Maas links clade. anatomy this of consisting in group a Nu- genera of identified 1.0. clearly studies never of have morphologically-based probability Olacaceae or posterior anatomically- a had merous but analysis 3-gene polytomy, a from emerging shows as tree species BI the MP three 3-gene the the shows Although tree monophyletic. as supported genus and well The cells is 1982). epidermal (Baas synapo- leaf pits in intervascular anatomical druses opposite two of presence and the data morphies: molecular ( by genera supported five com- 1 remaining clade Strombosia The and the C. clade woody of in crustaceous is posed and thin B is clade it whereas in the of fruit endocarp The drupaceous staminodes. flow- or isostemonous scales intercalary or with diplostemonous ers staminodes, has or C clade scales whereas intercalary without diplostemonous) (except flowers pentamerous tetramerous isostemonous or share subfamily this of Members eensis ld scmoe ftregenera, three of composed is C Clade ld scmoe fsxgnr with genera six of composed is B Clade Ptychopetalum and and , elspotda itrt h eane fteclade. the of remainder the to sister as well-supported Maburea , and , Strombosiopsis, Phanerodiscus eenvrcerbcueo t ihydivergent highly its of because clear never were Strombosiopsis Maburea Strombosiopsis, .grandifolia S. hscaercie 6 Sspoti the in support BS 66% received clade This . ,Atnra,Aaooee( Anacoloseae Aptandreae, ), to é and , o ta.20) ohmlclr(BI molecular Both 2004). al. et cot ,adAaooee( Anacoloseae and ), Heisteria and and ssse to sister as Tetrastylidium a elspotdwt mo- with supported well was Tetrastylidium Tetrastylidium Diogoa hra ayooy(Lo- palynology whereas Diogoa .philippinensis S. rtrplm Heis- Erythropalum, crdcru born- Scorodocarpus , Erythropalum Engomegoma .Teeatrela- exact The ). hr hyare they where hra these whereas , , sas well also is ) lx Dulacia Olax, ncls,Ca- Anacolosa, – Engomegoma mthick mm 2 iga En- Diogoa, Strombosia é o tal. et cot , nits in The . é Tet- cot , , Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 eea ihalrltosiswl upre.Tefrtclade first three The with supported. each contains well subclades, relationships mo- two all yield with The 3) genera, oblate). 2, or (Figs. (breviaxal trees lecular diameter shorter equatorial is axis in- the polar alternate than whose grains strands), pollen additional and cylinder pits, distal complex tervascular complete less style, a or cylindrical not more long but with (simple the all bundle are by vascular clade shared petiole be features this to Other of appears clade. ovary members the the for in synapomorphy locules sup- two a BS of low presence had The latter 1.0) port. the = although PP is data, 100, genera morphological = six (BS and of molecular group with monophyletic This as diver- evolution. resolved fruit remarkable and a flower by in characterized sity is and Santalales long basal a and petiole the of base style. the conical at incom- cylinder an F: vascular clade plete characterize synapomorphies two least At data morphological with support BS 90% and (Mal 1.0) = PP 100, (Mal endocarps or crustaceous staminodes thin with ( isostemonous pentamer- scales diplostemonous sometimes bear flowers, C Sleumer ous clade 1910; of Members (Gagnepain 1935). Erythropalaceae family, own a nlssbttemrhlgclcaitcsuyb Mal by study cladistic morphological the accres- but an analysis lar and dehiscence, genus The anther calyx. cent valvate petals, thin clade, long, second The pollen. with diploporate an- and with prolonged connectives, dehiscence, petals ther anther cells, porose thickenings, guard apical lignified synapomorphies: several with clade corresponding Mal the by analysis cladistic were logical data molecular No place to stamens. available of cycles two with flowers 63%, sup- = weakly (BS and data 1.0), morphological = of Mal PP analyses 100, on = based (BS ported data molecular of ses unclear. remains Cou- Santalales of basal placement among the leae diagnosability, clear its (Mal cell Despite flowers 2004). diplostemo- epidermal pentamerous polystemonous and leaves, to inflorescences, the nous thyrsoid in pres- spike-like channels walls, druses, laticiferous secondary cell of den- epidermal ence leaf lignified of presence hairs, the dritic as marked such is synapomorphies clade several The by Couleae. tribe and b), as 1961), circumscribed 1899a, Stauffer been 1935; (Tieghem has (Sleumer 1899 workers as subsequent all early by as recognized was genera aueo h ise novdvre.Freape nboth in exact the example, but For G, varies. clade involved in tis- tissues Anacolosa rule floral the the of be of Accrescence to clade. nature seems this fruits in on it sues placed (2004) al. et some to in (reduced flowers cyl- inflorescences vascular solitary complete umbellate vascu- a fundamentally petiole not ana- but inder), basal wood bundle, share using simple also (a (1984) larization taxa Oever These den features. van tomical by confirmed to taxon was de- this recently linked is (1980) most clade Lee The and this resolved. of not member in scribed Santalales in F clade 08 MAL 2008] ld ersnsoeo h otuuulgop of groups unusual most the of one represents G Clade = (BS data molecular from support strong received E Clade ld swl upre smnpyei ae nanaly- on based monophyletic as supported well is F Clade é é o ta.20) swt ld ,teeatpsto of position exact the E, clade with As 2004). al. et cot huohtn Aptandra Chaunochiton, o ta.20) hsvr ooeeu ru fthree of group homogeneous very This 2004). al. et cot Erythropalum ahda Anacolosa, Cathedra, and Cathedra Douradoa and Harmandia h lrlds sacecn n sur- and accrescent is disc floral the , esei asplundii Heisteria Ximenia ncaeF oee,temorpho- the however, F, clade in and , and Curupira é a nvial o molecu- for unavailable was é Ongokea, aai oleifera Malania o ta.(04 lcdi in it placed (2004) al. et cot o ta.2004). al. et cot Phanerodiscus, pce) n tetramerous and species), Ximenia , Malania É O IKET LCCA HLGN 103 PHYLOGENY OLACACEAE NICKRENT: & COT scaatrzdby characterized is hsrelationship This . n riswith fruits and ) and , hc have which rmChina from é o tal. et cot Ximenia é cot . ncnrs otemrhlgclcaitcaayi fMal of stands analysis cladistic position morphological This the 3), support. to (Fig. no contrast essentially BI in with With but J. position J group and clade sister I unsupported clades an with in taxon Mal this and includes (Nickrent data lar hl,w eomn nldn h 3seisof species 13 the including recommend we (Mal phyly, genus the within the case, genus this In 3). taxon (Fig. World BI New with obtained is resolution greater morphological with recovered of composed also Olax, clade, This was support. without and albeit 1.0) data, = PP 100, J. clade in in those sepals of loss in result or Octoknema tribe reduction a a it The made Opiliaceae), Schoepfioideae. who subfamily (1964) (near Schultze-Motel of order de- that to the more similar within a position indicated rived pollen 1982) of Studies (Lobreau-Callen 1984). other and Oever ultrastructure from den leaf van isolated 1982; on was (Baas work taxon Olacaceae this further indicated whereas anatomy 1955), wood wood (Mild- Reed Couleae and 1935; with trichomes relationship bread various close stellate to a suggested floral links anatomy example, conflicting For provide groups. or en- and laminae), ex- flowers, female pubescence, docarp the either stellate in are excrescences dioecy, features stigmatic (e.g. panded Morphological E. Santalales and for C, equivalent unusual B, taxa with clades placed our was genus to the where (2004) al. et ad14;Tktjn19)wihwstaserdt Santa- to transferred (1957) was Stauffer by which laceae 1997) Takhtajan 1948; nard their indicating thus 3), (Fig. tree evolution. BI molecular the of rates on those higher than taxa longer other comparatively are of genera branches these the Moreover, to pollen. leading the and on petals, apocolpium and concave fila- stamens a staminal the fused between tissue as glandular such ments, synapomorphies genera of number The a by evolution. gene homeotic the Aptandra from floral study to of candidate excellent membranous, perspective an into be would matures family variation, the this eventually Given vesicles. and lobed Mal laterally 2003) 1968; pseudo-inflated, al. (Capuron disc et the and cot calyx the between velops In maturity. induviale at drupe the rounds oNwCldna h genus of the members Caledonia, New African to to taxon The this stamens. section three links ovary and feature superior staminodes five vs. latter of half-inferior presence its the on and based 1886) (Valeton Olax of species two the for polytomy share a gives genera 2) three char- palynological of (Mal and acters group morphological, This anatomical, (1896). several Tieghem van by t w aiyb a ige 10) nsbeun works subsequent genus In the (1905). included often Tieghem family van in the placed by was family genus own The Africa. its tropical from trees of species B 0,P .)adrltosiswti hscaeare clade this within relationships and 1.0) = PP 100, = (BS revision. nomic ld a togyspotdb oeua aa(S= (BS data molecular by supported strongly was I Clade Octoknema ld srsle smnpyei ihhg support high with monophyletic as resolved is J Clade hsgnswsoiial eadda eto of section a as regarded originally was genus This . and Olax Triandreae and ” Ptychopetalum, tutr beti h lwra nhss de- anthesis, at flower the in absent structure a , si gemn ihohrsnaaentx uhas such taxa santalalean other with agreement in is é oiinmitie ihicesdsampling increased with maintained position a , o ta.20) lhuhte3gn Pte (Fig. tree MP 3-gene the Although 2004). al. et cot h oemme fCaeH sagnso 6 of genus a is H, Clade of member sole the , Ongokea ihc.4 pce itiue rmAfrica from distributed species 42 ca. With . Dulacia r lal lsl eae sevidenced as related closely clearly are é o 02.Tu,t ananmono- maintain to Thus, 2002). cot – a eonzda lcca .str. s. Olacaceae as recognized was sebde ihnteOdWorld Old the within embedded is lcmn upre ymolecu- by supported placement a é o 01.TeM re(i.2) (Fig. tree MP The 2001). cot Olax Phanerodiscus eursamdr taxo- modern a requires Octoknema Olax LusadL and (Louis the , and ssse to sister is Dulacia Dulacia, Dulacia “ coupe Olax Olax é é cot – o- in é 7 - , Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 s.Gvnti hlgntcifrain n a recon- may one information, phylogenetic parasit- this by Given conferred advantage ism. considered selective the are given condition the unlikely autotrophic in the clades to remaining hemiparasitism Mal polytomy the in to argued the As sister of order. is resolution Ximenieae that require such would This (i.e. parasites. Ximenieae all that hemipara- Malania root predict of can origin one single sitism, a assuming and trees, lecular and F G parasitism clade on clade which exists for information from MP no polytomy Moreover, the emerge. the E neither and resolved Unfortunately, analyses of F. BI evidence clade nor positive be is which can exists for it parasitism clade E, including first (i.e. clade The these genera nonparasitic. for in three exist all reports not that the do assumed given members parasitic Moreover, C, that and clades. likely B is clades it in been occur thus to autotrophic) shown have been (hence not have nonparasitic that may be genera nature five para- the in of their studies occur Four thus documented. would these pots, as in because grown behavior is sitic This with out haustoria. presence carried root the were of of confirmation absence allow or not do (1961) Rodrigues Ongokea enotie o ieoaaeu genera: bosia olacaceous five 1969), for has obtained haustoria) of been absence documented (i.e. nonparasitism for and chopetalum for documented genera: been four has parasitism only root actual lat., s. caceae distinct data as these 33 Schoepfiaceae of [Volume Olacaceae. of All from recognition 1955). olaca- all support (Reed of strongly derived examined most the genera was ceous ratio length trac- possesses vessel its it to and that heid 1984b) in (Sleumer Olacaceae parenchyma from aliform-confluent distinct also is anatomy that stated (1894) Engler calyculus. epfia anatomy, a 99, wood called of structure basis = vestigial the a On (BS to reduced support calyx a with strong has As shows 1.0). = press) PP (in Vidal-Russell of Nickrent study gene and five the relationship but this support, genes, Misodendra- low three receives to using study, sister present as the Mal In taxon and ceae. this Nickrent placed 1998; consistently al. have et Nickrent BOTANY 1996; and SYSTEMATIC Duff (Nickrent analyses molecular Previous Olacaceae. except 2004), 1896; al. (Mal (Tieghem analyses et family cladistic cot own morphological its In 1910). to Ola- Gagnepain and dis- in subfamily 1894), As or (Engler J. tribe caceae a clade 1830), in (Candolle Loranthaceae occurs to that Mal lat. in s. cussed Olacaceae of member Mal and (Nickrent phy- Santalales molecular of previous studies in shown logenetic to been are has they than This Viscaceae, Olacaceae. this and Santalaceae and members e.g. derived 1.0) Santalales, PP. more of 1.0 to = and related PP support closely BS more 99, 100% are Opiliaceae with = Opiliaceae (BS to sister Loranthaceae is clade to Schoepfiaceae/Misodendraceae sister is The clade resolved. fully also 104 n a eonzdb aeo (1886). Valeton by recognized was and h vlto fPrsts nSantalales in Parasitism of Evolution The Ximenia Pn 97.Suiso h emnto of germination the of Studies 1997). (Ping a the was rtrplm caotcy,Scorodocarpus Ochanostachys, Erythropalum, ,a ela l padeeand Aptandreae all as well as ), yHce 10,10)a ela hs of those as well as 1902) (1901, Heckel by Aslio1932), (Anselmino Hce 90 abr10b.Pstv evidence Positive 1907b). Barber 1900; (Heckel “ é oteovdgnso Olacaceae. of genus evolved most o ta.(04,ti eu a enassigned been has genus this (2004), al. et cot Schoepfia Octoknema Olax Misodendrum Bre 97;Pt ta.1990), al. et Pate 1907a; (Barber é o ta.(04,rvrasfo root from reversals (2004), al. et cot a itrt l atllsfamilies Santalales all to sister was ie h oooyo h mo- the of topology the Given . Schoepfia n Loranthaceae, and Wrhe l 1979), al. et (Werth Schoepfia Octoknema Heisteria Minquartia — Strombosia Curupira mn Ola- Among and , Curupira ” stesole the is é é o 2001) cot o 2001) cot t wood Its Schoepfia r root are Strom- (Kuijt Scho- are ) Pty- and and by é - lcca ihrslsfo hsmlclrpyoeei analysis phylogenetic molecular this from ( results with Olacaceae rdtoa classification Traditional utpsto fgnr fe ihu lc [...] with place without sequence, often by more genera family but of them, a juxtaposition between even a affinity some not having is tribes, make it distinct which characters, family; genera sexual natural between the affinities a in the nor discover one characters, does external the in ther Olacac Plantarum, who (1910), Gagnepain French): families. smaller Tieghem van all of accepted by number accepted it a divided been who into b) not (1896,1897a, has Tieghem particular unit in workers, phylogenetic single a are day present humid the of does margins type the as ancestral as savannas, such This and areas forests fruits. dryer large in relatively occurred likely with tree being small as ancestor a parasite root santalalean ancestral the struct e ol Semr18a lcca.SneSleumer Since Olacaceae. 1984a) and (Sleumer even 1980) World (Sleumer New intact, of Asian/Malesian essentially of classification revision remained his The following that of 2). system modification (Table a Englerian was the family (1935) Sleumer or by proposed tribe Olacaceae at of them rank classified who the (1894) Engler by diagnosed those with Brachynema T lsiiaino Olacaceae of Classification ayo h ldsdsrbdi h rsn okcorrelate work present the in described clades the of Many ABLE uiu affinities Dubious Engl. Schoepfioideae Subfamily Sond. Olacoideae Subfamily Airy-Shaw Anacolosoideae Subfamily lue 18b,adBeee ta.(1996) al. et Breteler and (1984b), Sleumer rb .Atnra Engl. Aptandreae 6. Tribe Horan. Olaceae 5. Tribe Engl. Ximenieae 4. Tribe Engl. Anacoloseae 3. Tribe Dumort. Heisterieae 2. Tribe Engl. Couleae 1. Tribe Octoknema Erythropalum Schoepfia Ongokea Harmandia Aptandra Ptychopetalum Olax Malania Dulacia Douradoa Curupira Ximenia Tetrastylidium Strombosiopsis Strombosia Scorodocarpus Phanerodiscus Engomegoma Diogoa Cathedra Anacolosa Heisteria Chaunochiton Ochanostachys Minquartia Maburea Coula .Cmaio fataiinlifaaiilcasfcto of classification infrafamilial traditional a of Comparison 2. “ sudrto yBnhmadHoe nGenera in Hooker and Bentham by understood As xldd e text). see excluded, .CaeI Clade L. al.CaeE Clade Baill. xl n ednaCaeB Clade Mendonca and Exell el ld I Clade Vell. hnadSKLeCaeF Clade S.K.Lee and Chun .CaeF Clade L. ireCaeG Clade Pierre asCaeC Clade Maas ir ld G Clade Miers aq ld C Clade Jacq. ..lc ld F Clade G.A.Black crbrCaeJ Clade Schreber ir ld G Clade Miers lue ld ? F Clade Sleumer Bue lm ld G Clade Blume (Blume) lm ld B Clade Blume ireCaeH Clade Pierre iree al.CaeG? G Clade Baill. ex Pierre ul ld E Clade Aubl. – rtlrCaeB Clade Breteler et.CaeG Clade Benth. lm ld C Clade Blume oiidfo nlr(1894), Engler from modified aaoCaeG Clade Cavaco é ec ld B Clade Becc. nl ld B Clade Engl. et.CaeI Clade Benth. nl ld B Clade Engl. at ld E Clade Mast. scntttsahtrgnosgop nei- group; heterogeneous a constitutes es ’ lsiiain rt tasae from (translated wrote classification, s — h ocp htOlacaceae that concept The Ximenia oeua clades Molecular ” seFg.2 3) 2, Figs. (see . ’ clas- s . Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.210 on: Fri, 24 Sep 2021 07:54:10 aee ru fpat hthv entxnmclycon- taxonomically century. been a have over that for Ola- plants troversial of of reclassification group a a towards caceae, step first of the understanding provided our and improved have These analyses data. phylogenetic molecular among additional relationships require will of Santalales resolution basal further thus these clades, among not seven previously relationships do the date all for to evidence obtained have unequivocal results yield that The Olacaceae. clades in placed additional been evi- seven provide for trees dence molecular the to Schoepfiaceae), addition as In classified paraphyletic. be to shown morphologically-based Mal were the of with analysis agreement cladistic in is result polyphyletic. This are Olacaceae analyses, phylogenetic molecular taxa of phy- place. empirical assemblage in an logeny heterogeneous without problems this classification real short, it posed of In modification 2). slight a (Table the or into classification data their Englerian incorporate existing to attempted authors the cases e,J n .L iket 08 oeua hlgn fSantalaceae 231 of Pp. Olacaceae. phylogeny 1894. molecular A. A Engler, 2008. Nickrent. L. D. and J. Der, 1968. R. Capuron, 1830. de. P. A. Candolle, h uhr r epygaeu oteidvdaslse nApni 1. Appendix in listed Pal individuals de the Laboratoire to the Therefore, of grateful rarely impossible. Members been deeply these have are from would authors material plants the collected genetic seldom assembling or world, seen the around from rtlr .J,P as .D oswne,F omn n .Lobreau- D. and Bouman, F. Boesewinkel, D. F. Baas, P. J., India. F. southern Breteler, of trees Parasitic 1907b. A. C. Barber, of haustorium The parasitism. root in Studies 1907a. A. C. Barber, Olacaceae, the of classification and anatomy Leaf 1982. P. Baas, Muira-puama. Droge der Stammpflanzen Die 1932. E. Anselmino, MAL Oever den and e.g. van 1992) named, al. 1982; been et have Baas genera 1955; new 1982) and (Reed 1980, 1984) anatomy Lobreau-Callen paly- from 1978; from Feuer e.g. and 1955; amassed, (Reed been has nology evidence new sification, 2008] ieCmu eBiosyst de Commun vice nlss n o rvdn nulse N eune fLorantha- of sequences DNA unpublished work, providing ceae, laboratory for with Duff, and assistance for analysis, Joel (SIUC) R. Vidal-Russell Romina Cabrera, thank Jonathan Garc Carbon- A. dis- assistance: University, and Miguel Illinois laboratory comments Southern provided helpful at for dale people thanked following warmly The also cussions. are Paris, in Curie a upre ygat rmteUS S:DB9094 MCB-9808752, DEB-9407984, NSF: here DEB-0108229. U.S. reported the and work from The grants sequencer. by DNA supported automated was her of use us lowing A srcgie yerywres n o ofre by confirmed now and workers, early by recognized As CKNOWLEDGMENTS (Santalales). Botanique. Section Mad car, de arborescentes ou arbustives gascar Santalacées et Opiliacées Olacacées, digesta W normas et naturalis Treuttel methodi juxta hu- cognitarum, plantarum specierumque cusque generum, ordinum, contracta enumeratio sive o.31,es .Ege n .Pat.Lizg ihl Engelmann. Wilhelm Leipzig: Prantl. K. and Engler A. eds. 3(1), vol. Gabon. from nus 1996. Callen. Society Philosophical Cambridge Botany Indian of Agriculture of Department 1 the of Memoirs scandens. and Berlin-Dahlem zu Museums und 626. Gartens Botanisches des blatt Misodendrum, – 47. rtrplm Allertonia Erythropalum. tnnrv:Cnr ehiu oete rpclMadagas- Tropical Forestier Technique Centre Atananarivo: . í ,EiaNcosn n akO Mark and Nicholson, Erica a, ytmtcBotany Systematic Engomegoma ü aéiu orléued afoefrsir eMadagascar: de forestière flore la de l’étude pour Matériaux rtz. Engomegoma and ihu h sitneo ueoscollectors numerous of assistance the Without . oaiceJhbce ü Systematik für Jahrbücher Botanische Schoepfia é L rdou ytmtsntrlsrgivegetabilis, regni naturalis systematis Prodromus aiu,bt tUniversit at both matique, ITERATURE é rtlr(lcca)anwmntpcge- monotypic new a (Olacaceae) Breteler o ta.(04 hr Olacaceae where (2004) al. et cot hnst .Spsfrgnrul al- generously for Sipes S. to Thanks . :155 3: – é Beee ta.19) u nall in But 1996). al. et (Breteler 3 npress. in 33: 4 246 14: 4 in 242 btnqee Pal et obotanique – C 210. ITED – i aülce Pflanzenfamilien natürlichen Die 256. É O IKET LCCA HLGN 105 PHYLOGENY OLACACEAE NICKRENT: & COT ’ el npriua,we particular, In Dell. é o é Maburea é Schoepfia rceig fthe of Proceedings ooi n Ser- and cologie iree Marie et Pierre 1:113 118: o.4 Paris: 4. vol. Octoknema 1 623 11: (Maas – (here Notiz- 132. Olax a- 2: – Mal Mal Lobreau-Callen, D. Hiepko, P. Boesewinkel, D. F. Baas, P. M., J. P. Maas, L J. and J. Louis, affinit et Structures 1982. D. Lobreau-Callen, iket .L,R .Df,A .Clel .D of,N .Yug .E. K. Young, D. N. Wolfe, D. A. Colwell, E. A. Duff, J. R. L., D. Nickrent, ora-aln .18.Caract 1980. D. Lobreau-Callen, iket .L n .Mal V. and L. D. Nickrent, plants parasitic of studies Molecular 1996. for Duff. J. methods R. sequencing and rapid L. D. film: Nickrent, to field From 1994. L. D. Nickrent, 42 Pp. Octoknemaceae. 1935. J. Mildbread, Mal Mal 1980. S.-K. Lee, 1969. J. Kuijt, uj,J 98 uulafnte fSnaaenfamilies. Santalalean of Dono- affinities J. Mutual M. 1968. and J. Kuijt, Stevens, F. P. Kellogg, A. E. Campbell, S. C. W., Judd, The 1997. Liang. H. and W. K. Hilu, usin approach an phylogenies: on limits Confidence 1985. J. Felsenstein, Beitr 1948. F. Fagerlind, 1924. Gilg. E. and A. Engler, ee,S .17.Aetr vlto ntegenus the in evolution Aperture 1978. M. S. Feuer, ekl .10.Srl emnto des germination la Sur 1902. E. Heckel, genres les dans germinatif processus le Sur 1901. E. Heckel, du parasitisme Amsterdam: le 1.7.1. Sur 1900. ver. E. 4Peaks, Heckel, 2006. Groothius. Corporation. T. Codes and Ola- Gene Arbor: A. des Ann Griekspoor, famille Sequencher. 2000. la Codes. comprendre Gene faut-il Comment 1910. F. Gagnepain, é é é é genre VI. (Santalales) of identity The Systematik 1992. für Welle. Jahrbücher ter Botanische H. J. Z B. and known Oever, den van L. Jahrbu Botanische Octoknemataceae. et Erythropalaceae Dipentodontaceae, terygaceae, Olacaceae. des et vltoaysuiso aaii lns p 211 Pp. plants. and phylogenetic parasitic Molecular of 1998. studies C. dePamphilis. evolutionary W. A. C. 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Acade :723 1: – é – :97 5: 470. ü r 168. é lce Pflanzenfami- rlichen 0 145 70: naso Botany of Annals é om yed type comme ´ so Icacinin ou es rbcL ytmtcBiology Systematic i e Sciences des mie – israinde Dissertation . – dnoi s Adansonia 100. 728. ’ and , ylbsder Syllabus s 3:381 133: – eis1vol. 1 Series (Olacaceae), monograph 154. 9 45 19: ’ é nseul un atpB Gardens te 0 29 10: New . ´ – é Bota- – ’ i 3 rie 461. une é Ok- 58. 19: 65: se- es. 1: – - , erl .. Asia, S.E., Merrill 2815 Nickrent DQ790169. DQ790130; L24079; DQ790189. Q917 DQ790166. DQ790127; Africa, AY042621. Hiepko, L26076; L24407; 2816; DQ790170. DQ790131; sylvestris Lepionurus GenBank and Nickrent) L. D. by accession rDNA, DNA SIUC for by at numbers followed archived are (samples SIUC) numbers at (deposited specimens barium 553 Mendon ot Rica, Costa rtlr qaoilGuinea, Equatorial Breteler, DQ790187. DQ790150; DQ790107; 4988; Q912 DQ790171. DQ790132; rubra lue,Peru, Sleumer, DQ790200. DQ790164; DQ790111; N/A. DQ790184. DQ790147; DQ790179. Rica, Costa Ducke, 6782 (Engl.) al. et ter DQ790178. DQ790141; Peru, Miers, eppig) 692 Sirikolo 2599 Zealand, DQ790172. New DQ790133; Schlecht., & Cham. f.) (Forst. s.n. Lamont 61 248 Q919 DQ790186. DQ790149; L24428; 2601; DQ790190. DQ790154; 3300 concinna Heisteria Guyana, DQ787438. gentina, Q918 DQ790194. DQ790158; pustulata Strombosia Philippines, DQ790192. DQ790156; DQ790123; DQ790195. borneensis 14745 Breteler E. DQ790180. DQ790143; DQ790122; 14888 DQ790173. DQ790136; emirnensis DQ790197. DQ790161; Mal Australia, 291 . DQ790183 DQ790146; amentacea nostachys Rica, DQ790188. Costa DQ790151; DQ790115; 4158; DQ790201. DQ790165; 497 al. et zinger DQ790199. Africa, W. Smith, et,C . .V ar,adL .Msemn 99 otprsts in parasitism Root 1979. Musselman. J. L. and Baird, V. W. R., C. aer Werth, of origins mistletoes: first The Nickrent. L. D. and R. Vidal-Russell, Olacaceae. the of anatomy wood Comparative 1984. L. Oever, den van gnnr macrocarpa Agonandra A 50 Q917 Q919 DQ790176. DQ790139; DQ790117; 4560; ; 25 Q919 Q917 DQ790174. DQ790137; DQ790109; 4245; ; é PPENDIX 59 248 125 DQ787447. L11205; L24418; 2599; ; 05 Q915 Q915 DQ790191. DQ790155; DQ790125; 4055; ; Schoepfia 177 Pp. aaiimi Santalales. in parasitism ference o,Sht .Bse,Madagascar, Bosser, J. & Schatz cot, 14 Q910 Q910 DQ790177. DQ790140; DQ790120; 4184; ; rtrplmscandens Erythropalum Srn.f)Ble .Africa, S. Balle, f.) (Spreng. ç .E rnsn,G mc 136 Amico G. s.n., Bran E. D. .F rvs 3796 Prevost F. M. ,Gabon, a, Biln ec,Malaysia, Becc., (Baillon) .Cae2176 Chase M. 27 Q914 Q914 DQ790181. DQ790144; DQ790104; 4247; ; d .Sd.Tuua skb University. Tsukuba Tsukuba: Sudo. S. ed. , ae,Madagascar, Baker, uai candida Dulacia tobsagrandifolia Strombosia aue trinervis Maburea ncls papuana Anacolosa 77 Q913 Q914 DQ787446. DQ790134; DQ790103; 2747; ; .Cleto nomto o N ape nlzd Her- analyzed. samples DNA for information Collection 1. .Ncrn 2758 Nickrent D. .Srets.n. Sargent S. – ol edulis Coula 7 in 178 85 204 Q918 DQ790167. DQ790128; L24084; 2815; ; .Mduys.n. Medbury S. 24 /;D704;DQ790182. DQ790145; N/A; 4244; ; crb (Olacaceae). Schreb. 22 Q913 Q912 DQ790198. DQ790162; DQ790113; 4232; ; .vndrWrf&R aqe 13875 Vasquez R. & Werff der van H. 22 Q911 Q918 DQ790175. DQ790138; DQ790121; 4212; ; .J eid .dWle11212 deWilde B. & deWilde J. J. .Frtetrs.n. Forstreuter W. .Cek58,M hs 843 Chase M. 5985, Cheek M. tnl,CsaRica, Costa Standl., at,Jv a Bogor), (at Java Mast., rceig ftePcfcRgoa odAaoyCon- Anatomy Wood Regional Pacific the of Proceedings .vndrWrf&Vsuz13846 Vasquez & Werff der van H. rbcL lvr Gabon, Oliver, .Weig 3288 Wieringa J. lm,Java, Blume, plaamentacea Opilia ahdaacuminata Cathedra noe gore Ongokea uti floribunda Nuytsia esei densifrons Heisteria aai oleifera Malania iei americana Ximenia tobsosstetrandra Strombosiopsis copi schreberi Schoepfia aaedo punctatum Gaiadendron Octoknema uuiatefeensis Curupira .O ilas ot Rica, Costa Williams, O. L. and , 18 Q918 Q915 AY042620. DQ790135; DQ790118; 4168; ; al. Gabon, Baill., 79 213 202 DQ787445. L26072; L24143; 2729; ; Pepg .Knz,Ecuador, Kuntze, O. (Poeppig) .Zmr 1928 Zamora N. .Sner 18-81 Senterre B. 78 236 Q918 DQ790185. DQ790148; L24396; 2758; ; lm,Jv a Bogor), (at Java Blume, as Guyana, Maas, 24 Q912 Q910 DQ790196. DQ790160; DQ790112; 4254; ; 81 F309 Q917 DQ790193. DQ790157; AF039079; 2831; ; ln ytmtc n Evolution and Systematics Plant celneg ooo Islands, Solomon Schellenberg, matK asealeptostachya Cansjera okrf,Gabon, f., Hooker .Sht ta.3620 al. et Schatz G. tcoeau petiolatum Ptychopetalum Biotropica p,Eutra Guinea, Equatorial sp., tobsaphilippinensis Strombosia .P e s.n. Teo P. S. 04 276 Q919 DQ790168. DQ790129; L24746; 3014; ; respectively. , Ha ire Gabon, Pierre, (Hua) etroaoii marquesii Pentarhopalopilia .Hmais.n. Hambali G. .Weig 2781 Wieringa J. esei cauliflora Heisteria .Senrs.n. Steiner K. hn&Le China, Lee, & Chun ob,Australia, Roxb., ioedu linearifolium Misodendrum .Agpre s.n. Augspurger C. 08 Q918 DQ790152; DQ790108; 3078; ; Lbl. .B. .Australia, W. Br., R. (Labill.) 89 51 237 L26074; L24397; 4591; 2829, ; .Weig 3295 Wieringa J. .Cae1329 Chase M. Bnh)Mes Brasil, Miers, (Benth.) igazenkeri Diogoa . Bahamas, L., mln Bahamas, Gmelin, nl,Fec Guyana, French Engl., lc,Brazil, Black, 02 Q916 DQ790142; DQ790106; 3052; ; iqataguianensis Minquartia 55 Q910 DQ790153; DQ790110; 4555; ; .Mlo 2575 Molloy B. .Zg s.n. Zagt R. .Sht ta.3439 al. et Schatz G. 1 140 11: 16 216 DQ790163; L24146; 4166; ; lxbenthamiana Olax 08 594 DQ790159; U59934; 3028; ; ersyiimperuvianum Tetrastylidium nl,Gabon, Engl., Ri.&Pv)G Don., G. Pav.) & (Ruiz. hnrdsu capuronii Phanerodiscus noeoagordonii Engomegoma .Ncrn 2764 Nickrent D. .Beee 15457 Breteler E. padatubicina Aptandra – huohtnkappleri Chaunochiton hmeeamanillana Champereia et. Australia, Benth., .Cae1328 Chase M. 143. 10 DQ790102; 4180; ; 89 DQ790101; 2879; ; 02 AF039078; 3042; ; 05 DQ790119; 4035; ; 04 DQ790124; 4054; ; 25 DQ790126; 4205; ; 22 DQ790105; 4202; ; 26 DQ790114; 4256; ; 17 DQ790116; 4167; ; esei parvifolia Heisteria .Ncrn 2601 Nickrent D. ueaantarctica Tupeia .Ncrn 2816 Nickrent D. .Ceets.n. Clement C. Eg. xl & Exell (Engl.) .J Mac J. M. 72 L24146; 2732; ; mt,French Smith, .Beee tal. et Breteler E. lvr Gabon, Oliver, 72 L24425; 2742; ; .Sner SO Senterre B. 09 N/A; 3079; ; Scorodocarpus Bil)Rolfe, (Baill.) amn 340 Caoming i press). (in .Nickrent D. Moquiniella eaao& Regalado .Wieringa J. .A Rat- A. J. C Ar- DC, Miquel, .Mun- J. (Engl.) 4204; ; í 4165; ; Aubl., 4268; ; 2764; ; aetal. Ocha- Olax (Po- ial D. B. ; ; ; ;