Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Taxonomists. All rights reserved. sa h eu setmtdt opiemr hn50spe- after 500 family than third-largest more the tropical it comprise making to in cies, estimated is aroid genus The understood Asia. well least and complex, O 10.1600/036364413X670430 DOI © Botany Systematic diin yFrao(99 n ot 16) aoe al. et Mayo (1967). sections: Hotta five and recognize (1939) with (1997) (1912), Furtado Engler and by (1860) Schott additions of work the upon based and ls(..iae,ifoecne nachl o suspected for alcohol) 2008). in Wong and inflorescences (Boyce novelties sam- images, adequate often prepare (e.g. is and it ples localities, available, key re-visit How- are to data pollination. possible locality during concise provided damaged ever, irreparably been have staminodes, time interpistillar which notably morphologies, by floral post-anthesis critical collected bee- specimens by (2) depredation indeter- and post-preservation tles, effectively (1) is incorrect to: material owing have minable the or of undetermined Much either determinations. are specimens of ity 01 ogadBye21;Wn ta.21) ept the Despite al. 2011). et al. Kurniawan et b; Wong al. 2011a, of 2011; al. abundance et Boyce et and Boyce Hoe Wong 2010; 2010; 2011; al. Boyce et and C. (Boyce Tung (P. Baharuddin described 2010; recently date 2008; are to Wong 17 available which and of are Guinea Borneo obs.), names pers. on New accepted Boyce, focus 30 currently and only Studies Borneo, where 2008). Wong Sumatra, and diversity: three diversity (Boyce are there of greatest where Asia the East and centers South with of forests species tropics, tropical the Asian of in and majority Neotropics overwhelming the circumscription, in current distributed the on Based 2011). oyih 03b h mrcnSceyo ln Taxonomists Plant of Society American the by 2013 Copyright Homalomena Homalomena opooia n hmclmre piiain osrce pteadfu tmn e tmnt lwraepeimrhcfor plesiomorphic of are subsequent species for Neotropical staminate used of per was separation stamens the taxa support four 15 and further and representing Homalomena that The accessions spathe supergroup. 20 constricted Punctulata of A and the phylogeny optimization. into reduced marker complex Homalomena. A chemical Havilandii supported. and and well complex morphological are Insignis to supergroups the assigned transferring Chamaecladon currently after species monophyletic Neotropical several is supergroups, Asian all of representing included; are taxa ie h sa pce) n uprsadrfnstegopn fAinseisit supergroups. into species Asian of grouping the refines and supports and from species), species Asian Neotropical the the (i.e. of removal supports study xeto of exception uegop.Aceia akra R at marker chemical A supergroups. .9mn n .0mnaesae mn h aoiyo aasmldfrAsian for sampled taxa of majority the among shared are min to 2.90 and min, 2.69 Abstract— Keywords— Philodendron 2 Homalomena rgDsoeyLbrtr,Cne eerhIiitvsFudto,1AJlnT ,TmnPrnutinUEP, Perindustrian Taman 5, TP Jalan 12A Foundation, Initiatives Research Cancer Laboratory, Discovery Drug .vivens H. hlgn fAsian of Phylogeny 1 21) 83:p.589–599 pp. 38(3): (2013), eateto ln cec n niomna clg,Fclyo eoreSineadTechnology, and Science Resource of Faculty Ecology, Environmental and Science Plant of Department ogSnYeng, Sin Wong Homalomena coti h otseisrc,taxonomically species-rich, most the is Schott hmclmresietfe rmlqi hoaorpyms pcrsoypoiigpoie nidpnetsto markers of set independent an provided profiling spectroscopy chromatography-mass liquid from identified markers Chemical . hlgntcaayi ftegenus the of analysis phylogenetic A oaoeavivens Homalomena tmndsaogtepsilt oeaels he ie neednl nHmlmn uegop ucuaasupergroup, Punctulata supergroup, Homalomena in independently times three lost are zone pistillate the among Staminodes a enpeiul iie nosections into divided previously been has 3 ono hrce apn,sseais . systematics, mapping, character Borneo, colO ilgclSine,Uiest an aasa 10 S,PluPnn,Malaysia. Pinang, Pulau USM, 11800 Malaysia, Sains Universiti Sciences, Biological Of School Asian . hwtouiu ek tR at peaks unique two show cot(oc ta.21;ByeadCroat and Boyce 2010; al. et (Boyce Schott Homalomena pcmn nhrai,temajor- the herbaria, in specimens nvriiMlyi aaa,930Smrhn aaa,Malaysia. Sarawak, Samarahan, 94300 Sarawak, Malaysia Universiti 1,4 hmclmre tR at marker chemical A . Curmeria a e Jean, Pei Tan swl upre smnpyei n xldsNeotropical excludes and monophyletic as supported well is 4 ihMrhlgcladCeia Data Chemical and Morphological with t Homalomena uhrfrcrepnec ([email protected]) correspondence for Author .0mni pmrhcfrteCaacao uegop ihasprt anin gain separate a with supergroup, Chamaecladon the for apomorphic is min 2.80 aiudnB Ahmad, B. Fasihuddin t Lne Andre & (Linden .5mnad35 i.Fv hmclmresspotteCroldnsprru ihthe with supergroup Cyrtocladon the support markers chemical Five min. 3.54 and min 3.25 70 uagJy,Slno,Malaysia. Selangor, Jaya, Subang 47600 omnctn dtr akP Simmons P. Mark Editor: Communicating Anthurium Homalomena Homalomena 2 gKa Kiaw, Kiaw Ng Homalomena t .0mni lsoopi o h hmeldn oaoeaadPunctulata and Homalomena Chamaecladon, the for plesiomorphic is min 3.60 Schott Aaee ae nteISRgo Combined Region ITS the on based () Aaee oaoeee ae ntenT eini rsne.Eighty-nine presented. is region nITS the on based Homalomeneae) (Araceae: Homalomena ´ is ) 589 eeie h opooia onaisof boundaries morphological the redefines , 1 inof tion and K.Krause, & Engl. Krause); Neotropics; (including the Krause K. & Engl. ByeadWn 08 ge l 01,tels en a being last the 2011), al. et Ng Hotta’s for 2008; name replacement Wong and Punctulata Wise Asian (Boyce and and of Cyrtocladon, sections Chamaecladon, Hay four Homalomena, 1998; The Hay 2000). 2001; Yuzammi Homalomena and Hay Hay and 1991; Boyce b; 2001a, intractable taxonomically Pothos other This in (e.g. undertaken. used groups is been testing has phylogenetic approach taxo- until Asian facilitate study to 2011), tools nomic Boyce useful as and complexes, and Wong supergroups and 2011; (Boyce al. papers et Homalomena previous Ng In 2008; history. Wong their in point some itlhih,ecpinlysaioe r bet( absent the are equalling staminodes staminode exceptionally associated height, the pistil three- usually with is locular, ovary four- the The anthesis to spathe. species at the from swiftly many protrudes elongates it spadix in until the although of move- portion spadix recorded, no staminate been and limb, have spathe ments the Spathe of spathe. closing simple and comprise upper gaping known, weak where and very anthesis, a lower during only movements the or no between with long, constriction tis- cm and 1.5 aromatic modules, exceeding shoot strongly spathes hapaxanthic with rarely or creeping pleionanthic sues, to erect large to n ee .Boyce C. Peter and h oaoeasprru S)cmrssmedium comprises (SG) supergroup Homalomena The rmteAintxn he hmclmresa R at markers chemical Three taxon. Asian the from 3 ha .Othman, S. Ahmad . and L., Geniculatae Homalomena Cyrtocladon Alocasia a iie noifra opoao units, morphotaxon informal into divided was eerdcdt nomlsprrus(SGs): supergroups informal to reduced were Rhaphidophora Homalomena n etoia pce urnl assigned currently species Neotropical and , l aebe eonsda eeaat genera as recognised been have all , Homalomena G.Don., 3 Gif)Furtado; (Griff.) Geniculatae 3 Homalomena e ogBoon, Hong Lee (‘ as. e oc 00,b c, b, 2000a, Boyce see Hassk.; Adelonema Geniculatae h ytcao supergroup Cyrtocladon The . Euhomalomena Homalomena .ot.Wt h excep- the With M.Hotta. n eetdspecies selected and , Chamaecladon .punctulata H. cot etitdto restricted Schott) . ol Moritzi, & Zoll. fEg.&K. & Engl. of ’ es stricto sensu 2 t .5min, 2.55 .expedita H. This . (Miq.) Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. xadd n h hcedhsigltrly Pistillate ovary laterally. the connective dehiscing staminodes; 4-locular. interpistillar the is thecae associated with lack the stamens and Staminate 3–4 constriction. expanded, comprise moderate or each weak are a flowers with long, the than cm more which plants, 2 spathe the in aromatic distichous, or modules sub-distichous medium-sized frequently shoot comprises pleionanthic 2011) with al. et Ng are connectives and stamens slightly laterally. dehiscing six) thecae have to very with expanded flowers five substantially Staminate even (rarely pistil. four or the to of three to, height the equal by, exceeded staminode with locular, associated four to three the are Ovaries anthesis. com- move- during a spadix ments and undergo spathe studied coordinated seemingly lower far of so series the plex species longer), between All constriction much spathe. pronounced often upper and to (and weak long modules, a cm shoot with studied) 2 exceeding been spathes have and species few very pleionanthic (albeit tissues, aromatic strongly with plants creeping to terminally. dehiscing the thecae and with unexpandedconnective, an locular, with stamens, Staminate three) three (seldom pistil. two have the to flowers than two shorter have much is movements staminode ovary associated spadix gaping No The limb. recorded. simple movement spathe been comprises the Spathe of known, spathe. closing and where upper constriction anthesis, and no with during lower long, than cm only 1.5 less the to known, up mostly between rarely is are very long, as Spathes cm 1 far rarely modules. very as shoot odorless, and pleionanthic with tissues, plants vegetative erect aromatic, often less creeping, laterally. a dehisce forming thecae connective the stamen; expanded each an over with cap stamens (rarely four six) to to three five have flowers Staminate Hersc.). & A.Hay negncsae oanalyse to spacer intergenic oee,dvriiaino h xatseisol started only species in extant 2012b). recently the al. comparatively et of Nauheimer diversification before 2004; years However, Bremer million (138 and old Janssen of is Araceae present; of species node stem Asian the that two only included (2008) DNA Homalomena al. ribosomal et two intron Gauthier chloroplast on the and based (ETS), spacer clade, scribed entire transcr internal the markers: nuclear to basal as on Homalomena focusing (2008), to and sister wallisii as H. (1997) al. et 9 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 590 i yCbeae l 20)placed (2008) al. et analy- Cabrera multi-gene a by However, sis polyphyletic. (1997) al. et Mayo Neotropical within two nested that result the hr rnhscaatrz h oeo h rw clade. region crown plastic evolving the slowly of the level, core intergeneric the the At characterize relatively and branches level, low intrageneric very short the is at There al. radiations. divergence et rapid sequence Low followed 2012a), 2010; then al. and al. et 2011), et (Wong Nakai (Nauheimer Schismatoglottideae Brongn. and (Schott) Colocasieae the Barabe SG; Geniculata as known (previously SG Punctulata The erect large very to medium comprises SG Cyrtocladon The often small, to minute comprises SG Chamaecladon The eetpyoeei okpoie opligevidence compelling provides work phylogenetic Recent .magna H. ´ ta.(02 sdteplastid the used (2002) al. et Hreun aslcino etoia species) Neotropical a of selection (a ‘Harlequin’ : .cochinchinensis H. ssse to sister as .a PpaNwGie).Guhe tal. et Gauthier Guinea)]. New (Papua A.Hay Philodendron Philodendron Philodendron Philodendron Philodendron Anthurium rendering , nl and Engl. bdsae IS,etra tran- external (ITS), spacer ibed Homalomena eovdNeotropical resolved , Homalomena trnL u Asian but cot(ale 2011), (Carlsen Schott and .philippinensis H. bsdo w taxa, two on [based nrnand intron Homalomena Homalomena rpl16 pce were species es Mayo sensu Homalomena However, . trnL–trnF Engl. sensu ,with ris(ilase l 91.Kt ta.(97 aeshown leaves have of in (HNJ) (1997) 2,5-dihydroxymethyl-3,4-dihydroxy- cc-homonojirimycin and al. of (DMDP) most et pyrrolidine of Kite presence distinguishing 1981). the properties al. that et irritating (Williams the aroids in involved 91 lad ta.20;H ta.20;Ta ta.2010). al. essential et reported Tian undetermined (2006) 2008; an al. al. for et et profiling Wong Hu region, oil al. Malesian 2004; et the al. (Zhou In et sesquiterpenoids Elbandy and 1991; monoterpenoids of other composed as of primarily identified plants oil of tial (Sung Rhizomes 2000). sesquiterpenoids al. occulta several et H. and Singh 1992; linalool al. 80% et oil to essential up an containing with rhizomes the with conta 1986), (Hegnauer oil essential identifications. 1.2% taxonomic yielded poor from aromatica medicinal purported (e.g. with application species on mainly focused patchy, studies. systematic in morpho- that data and logical illustrates the molecular above complement The with can (2008). characters congruent chemical al. et is Cabrera of which phylogeny Aglaonemateae), (tribe Engl. Nephthytideae), (tribe nsudsseai nesadn rrs en uieo even or futile being risk or understanding systematic basis sound a in requires an research pharmaceutical Asia, yet in activity tropical of and in increase to genus species-rich need aroid abundant, mesophytic a understood define most least from to the Aside study understand phylogeny. rigorous better their of resolve need and critical species in group complex as Hue annotated Schott from that (probably material type the the occulta and H. lost, as is applied Vietnam) anti-microbial cautiously in identified be that to 2010) name is al. the However, et activities. Tian anti-inflammatory and 2008; al. et Hu on out carried ‘ as httetotpso cdccfecai eiaie are derivatives acid caffeic acidic of types (80%), two probable highly Colocasioideae the carboxylic is It subfamilies that free (20%). Pothoideae in a and (38%) common Lasioideae with species), are derivatives non- group acid while of (20%) caffeic (67% are Pothoideae sulphated acid and Monsteroideae caffeic (23%) of subfamilies Philodendroideae esters of in Sulphates present 1981). C-glycofavones; are al. and others procyanidins et while (e.g. Williams acid), family the sulphuric to of unique esters taxa of investigated the sulphates to 1997). (e.g. unique are (Hegnauer compounds these raphides of oxalate Some calcium and cyanogenic flavonoids, glucosides, including sapo- compounds mainly are phenolic metabolites over- nins, sampled secondary However, aroid species. 1986) that seems known it the Hegnauer all of 1981; percentage tiny al. a only et Hottarum (Williams the in chemistry level taxonomic showed low (2011) a al. Schismatoglottideae). et at (Araceae: Low useful group (2011). Gauthier be al. data), to et Low (unpubl. ITS and al. (2008), et al. Buzgo et are intertribal to Exceptions taxonomic work. low-level Araceae for on region studies ITS appropri- phylogenetic the few more sampling very are are there spacers far So nuclear ate. evolving fast level, cific matK ulse hmsr eerhon research chemistry Published h w ulse opeesv conso Araceae of accounts comprehensive published two The Objectives— Nephthytis .sagittifolia H. ssial oifrrltosis hl tteinterspe- the at while relationships, infer to suitable is Srn. cot,adsfesfo rbesresulting problems from suffers and Schott), (Spreng.) ossso tlattreseis(oc n i2010). Li and (Boyce species three least at of consists eerpre oyedvroscmonso essen- of compounds various yield to reported were Schott, .occulta H. Homalomena oaoeaocculta Homalomena ) eea hraooysuishv been have studies pharmacology Several ’). Anchomanes Aglaonema and salreadtaxonomically and large a is .aromatica H. nn anymonoterpenoids mainly ining iaoladlwramounts lower and linalool Schott, Homalomena Lu. Schott, (Lour.) Homalomena cotand Schott oaoeaaromatica Homalomena Pseudohydrosme Wn ta.2007; al. et (Wang (misidentified ssomewhat is Homalomena Aglaodorum .occulta H. Engl. ˆ ´ Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 03 OGE L:PYOEYO SA OAOEA(RCA)591 (ARACEAE) HOMALOMENA ASIAN OF PHYLOGENY AL.: ET WONG 2013] atire l 20)wr bandfo eBn.Frytotx of taxa Forty-two GenBank. from obtained were (2008) Asian al. 1). et from (Appendix sequences Gauthier Forty-five genes GenBank rRNA into JQ955571–JX076808. numbers deposited nuclear Accession and the under generated of newly were region sequences ITS2 Forty-four and 5.8S, ITS1, of . MMgCl mM 2.0 bnpol ftearea. the of people indigenous thoseIban the sig- among are plants to medicinal species moderate as importance having selected nificant as The (2002) Christensen Sarawak. by and highlighted Batang Aman, from mainly Sri sampled part Ai, taxa third for spectros- profiling chromatography-mass The (LCMS) liquid copy supergroups. involves delimit study the the to on of focuses used that study currently characters the morphological are of eight part of second patterns The evolutionary 2011). al. et 2008; Ng Wong and (Boyce supergroups proposed phy- the of testable topology a produce to for logeny are objectives Other reason. of same Species con- their status. test generic to and are included are supergroups Cyrtocladon, species Neotropical Asian Punctulata. Chamaecladon, the Homalomena, all for included: diver- taxa of centers Representative main the sity. of one of Borneo, Malaysian phylogeny from taxa the resolving on focuses Homalomena study This harmful. rmNnaSma aagA,Srwkbten20 n 2010. and 2009 between Sarawak Ai, Batang outgroup, Sumpa, An Nanga from Except 1). (Appendix study S13375). number (study TreeBASE to combined The deposited 1. been GenBank Appendix has in and matrix provided data information are taxa Voucher all for available. numbers ITS accession not al. as Callopsideae are (2008) et (Culcasieae, Anubiadeae) al. Homalomeneae (Wong et and of Cabrera tribes Alliance on sister based from Schismatoglottid selected sequences were and the outgroups from P.C.Boyce The 2010). selected & were ( S.Y.Wong Ridl.) Jonker) taxa & outgroup Jonker Two represented. are ecin PR)wr odce nattlrato oueo 20 of volume reaction chain total Polymerase a 1990). in 1 al. conducted et comprising were pairs (White primer (PCRs) respectively the reactions using 3F/4R, amplified were and al. ITS2 et 1F/1R and Wong subunit (Polyvinylpyrrolidone; 5.8S PVP ITS1, of 2010). addition the with 1987) Doyle ihtemlil-reoto nefc,adsvn pt 000tesfrom trees 10,000 to up addition. saving sequence branches, and random effect, zero-length each in of option multiple-tree collapse the of swapping, with searches bisection- branch tree heuristic addition, (TBR) sequences with reconnection stepwise obtained random with were replicates trees 1,000 parsimonious most The weighted. equally characters Phyloge- all with reconstruction data. (MP) parsimony max- similarity missing imum for 2002) following (Swofford as PAUP*4.0b10 with adjustment treated performed were manual analyses were netic Indels minor v5.6.4 2004). by Pro (Simmons followed criterion Geneious 2012) in from al. Drummond implemented et www.geneious.com; sequences Zealand; aligned as New was Auckland, with matrix 2004) Ltd., data (Biomatters The (Edgar combined (2010). al. MUSCLE et were Wong using for and sequences (2008) assembled al. and et These trimmed Gauthier manually taxon. were region each ITS the of sequences igecerbn a rsn h rdcswr etfrsequencing Bhd., Sdn. for Laboratories sent a BASE were if First at and products Malaysia. directions gel, Selangor, the reverse agarose present and 1% Lithuania). forward was a in and Vilnius, band using gels, (Fermentas, clear again agarose kit viewed single 2.0% were purification or products PCR 1.5% Purified a on visualized using conditions. were purified cycling products standard PCR using amplified The were fragments ITS cation. n otwo to One xrce sn oiidvrino h 2 the of version modified a using extracted tissues. vegetative aromatic with Schismatoglottideae tribe of ao Sampling— Taxon euneAinetadPyoeei Analyses— Phylogenetic and Alignment Sequence N xrcin C mlfcto,adSequencing— and Amplification, PCR Extraction, DNA wnyacsin ersnig1 aawr nlddfrteLCMS the for included were taxa 15 representing accessions Twenty Homalomena m Homalomena fDS a de oec ecint mrv amplifi- improve to reaction each to added was DMSO of l

2 + cimtgotsnervosa Schismatoglottis units 2 , nldn eiyn t oohl,fcsn on focusing monophyly, its verifying including ufr . MdT i,02m fec primer, each of mM 0.2 mix, dNTP mM 0.1 buffer, i fNeotropical of six , aeil n Methods and Materials ihynn aawr nlddfrteanalyses the for included were taxa Eighty-nine Taq orsletevrct fadinternal and of veracity the resolve to , .expedita H. N oyeaead2 and polymerase DNA Philodendron Homalomena a eetda representative a as selected was , and hlnto americanum Philonotion

+ wallisii H. TBpooo Dyeand (Doyle protocol CTAB r nlddfrthe for included are cimtgotsnervosa Schismatoglottis n 9of 39 and , m l eecollected were all , el generated Newly fDAextract. DNA of l oa N was DNA Total Philodendron (A.M.E. m l 6 n 0 p n h egho T2vre between varied ITS2 of length the and bp, between varied 406 were ITS1 of products and length The PCR 264 band. single all a polymorphisms; as without resolved signal of unambiguous indication and clear any a provide region ITS nAcquity an (LCMS)— T eint sestepyoeei au ftemrhlgcland morphological the of the value on traced phylogenetic based the were analysis characters. assess chemical RAxML matrices to from The region tree ITS likelihood 1971). maximum Fitch 2010) the Maddison onto (unordered; and (Maddison parsimony 2.7.4 ver. using Mesquite matri- in charac- Character analysed non-ambiguous. all were are ces with characters All data binary. categorical as treated of presence/absence ters comprised as scored were four and Matrices = above, = characters. 1 described overtopping; 1 profiling not from stamens; chemical identified = shorter; the were two 0 markers = stamen: chemical = twelve 1 per Additionally, 0 connective overtopping. exceeding; flower: of or Shape staminate (viii) equaling per com- stamens; = in number staminode 0 staminodes): Stamen interpistillar pistil: of (vii) (Interpistillar Height associated (vi) zone with presence; = parison pistillate 1 absence; the = 0 among con- Staminodes Spathe ratio: length (ii) (v) spathe presence; Lower/upper characters: = (iii) presence; 1 = reproductive 0= absence; 1 = absence; seven = 0 0 lobes: and striction: posterior blade vegetative one (i) were states for character morphological Ancestral reconstructed 2011). al. et Ng 2008; Wong xrcswr re nvcoadtedidcueetat eekept were extracts crude dried the and methanol vacuo was combined 0 in procedure The at This dried extraction. agitator The temperature. total were room an powder. ensure ambient extracts on to fine at times methanol into hr three in 48 ground repeated for soaking and rpm by d 110 extracted at 10 was ca. material for ground air-dried were blades 50% construct to used were values trees. trees consensus with Remaining majority-rule index burn-in. as convergence discarded stan- as were the frequencies using out split by one < assessed of sampling was deviation and Convergence and trees dard generations. The cold random 100 one convergence. from every with starting of parameter generations chains, 2,000,000 assure heated for three to run Carlo was twice Monte algorithm MrBayes chain MCMC repeated Markov with was 2001). performed Ronquist (MCMC) and repli- were 10,000 (Huelsenbeck (Stamatakis running analyses 3.1.2 7.2.6 by ver. phylogenetic obtained RAxML were Bayesian using values bootstrap cates. out likeli- ML Maximum carried 2008). selected. al. were model et analyses substitution nucleotide (ML) the + hood was (GTR reversible G) time + Akaike General using I 1974). 2008) Akaike (Posada (AIC; 0.1.1 criterion ver. information jModeltest in selected were regions lcrsryinzto EI ore h orecplayvlaeand voltage 100 and capillary kV 2.70 source to set The an were temperature source. mass with source equipped (ESI) definition MA) ionization high Milford, electrospray Corporation, Synapt (Waters a detector to time-of-flight (oaTOF) acceleration coupled quadrupole-orthogonal (HDMS) system spectrometry (PDA) array diode ntoeue odlmtt h uegop of supergroups the delimitate to used those on pcrmty(RS nlsswscnimdwt tnadrun 2.1 C18, standard BEH Acquity a an Separation ppm). with on 1.5 obtained 195.0879, confirmed was measured 195.0882; per- was mass (calculated resolution caffeine was analysis of high spectrometer of (HRMS) Accuracy the Tuning M). of spectrometry (0.5 L/h. formate 700 calibration sodium calcu- at using pg/mL; while formed flow (500 556.2771) leucine-enkaphaline gas using [M+1]: dessolvation out lated and carried was V MS 6.0 the of to set was energy – i,1–0 rm5 i,adte eda %Ao 100% or dissolved A were extracts 0% 3 crude and The at filtered analysis mg/mL), The mode. (10 held min. ESI methanol 2 condi- positive in then for before in equilibrate min out and to 4 allowed carried min, for and was A acid) from 6 100% formic 5– A to 0.1% back 50–15% with from tioning min, (acetonitrile A 3–4 B from solution 0% with A of 15– 0.1% eluting 80–50% with min, min, (water mL/min 3 A in 0.5 4–5 solution A of 80% of 100% to rate of acid) formic system flow solvent constant gradient binary a a with column, UPLC .0 nepee sidctn odcnegne h is 0 ftrees of 10% first The convergence. good indicating as interpreted 0.005 hmclPoiiguigLqi hoaorpyMs Spectroscopy Chromatography-Mass Liquid using Profiling Chemical rprto fEtat o hmclProfiling— Chemical for Extracts of Preparation hrce Mapping— Character arxCharacteristics— Matrix h otsial uloiesbttto oe o aho h gene the of each for model substitution nucleotide suitable most The  ro oanalysis. to prior C < ;1=>1 i)Sai neto:0=srih;1=oblique; = 1 straight; = 0 insertion: Spadix (iv) 1; > = 1 1; CSaayi ftecueetat a efre using performed was extracts crude the of analysis LCMS TM lr efrac iudcrmtgah (UPLC)-photo chromatography liquid performance ultra eut n Discussion and Results opooia hrceswr eetdbased selected were characters Morphological h eunepeorm o the for pherograms sequence The m netdot h PCset-up. UPLC the onto injected L + 0m,1.7 mm, 50  ,rsetvl.Collision respectively. C, Homalomena h eil n leaf and petiole The m atcesize particle m Byeand (Boyce Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. h hmeldnS spol upre (ML supported poorly is SG Chamaecladon with The clade a 2011). forms al. Ar3047 et sp. Ng (sensu complex with Selaburensis clade the same the in supported strongly ee(i.2). morphological in (Fig. here complexes these the and Of delimit supergroups which tissues. characters morphological aromatic eight characters, of flowers, absence presence fertile and and anthesis, and during sterile movements the spadix and of spathe morphology the sizes, and in (Corner (ML Pocket supported Riau strongly However, 1.00). is the PP complex from 98%; Giamensis relicts Hanneae The The be 1960). complex. Hanneae might separated the taxa complex of these with remainder 1.00), the PP from from 96%, complex (ML Hanneae Sarawak the West with together placed and 91%, is Interest- Malaysia (ML complex. complex Rostrata increased the of support Rostrata ingly, exclusion the although with the 0.85) 1.00) PP PP of 61%, accession (ML single clandestina inclusion H. the the (with complex Hanneae of the it, Within 1.00). PP the of into absence fall the 1.00). 2011) PP supported 76%, moderately by (ML al. is supported clade et This placement staminodes. interpistillar (Ng a SG SG, Punctulata Cyrtocladon 1.00). SG Punctulata PP in previously complexes, 94%, The placed Insignis 1.00). (ML and Havilandii monophyletic PP the as However, 84%, supported (ML strongly group is a Chamaecladon and form Homalomena SGs the Together, shown). not lxi togyspotda aa oCroldnS (ML SG Cyrtocladon to the basal to 1.00). as sister PP supported 86%, com- is Borneensis strongly The complex, is 1.00). PP plex Hanneae 75%, (ML the complex Giamensis in placed tatively 0.85). Asian PP 64%, (ML the with monophyletic Within is SG Erythropus. Homalomena and Crinipes Neotropical Pte.). P. 9 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 592 scaeNWHi i.1 smnpyei n itrto sister and monophyletic is 1) Fig. in Philodendron N.W.H genus clade the Neotropical 1). as to Fig. 1.00; sister PP 84%, and (ML polyphyletic is (1997) presented is and tree S1). 0.57 consensus Fig. = data, (50% supplementary (CI) 0.78 in Index = Consistency (RI) steps, Index 2,718 Retention of parsi- length trees 642 a 100,000 constant, to of and limited were were searches uninformative, Tree characters informative. parsimony mony 541 but and variable 5.8S, aligned: ITS1, 331 were (partial positions ITS2) nucleotide partial 1,517 of total A olw aoe l 19,p8 i.6.Tepeec fleaf- of presence The in 6). plesiomorphic Fig. is p.8, lobes posterior (1997, blade al. et Mayo follows 2 n 9 p hl h egho h .Sgn pwas bp gene 5.8S the the of all length the among while bp, consistent 395 and 326 ab dpiet hohts,tepiayeooyo spe- of ecology there primary (and posterior the rheophytism, SG of to Chamaecladon adaptive absence the and maybe for The SG) plesiomorphic SG. (Chamaecladon is Homalomena lobes once in lost once or gained SGs, Punctulata Chamaecladon and in twice lost independently either are lobes rior .L 1. 86%, (ML supported strongly is SG Cyrtocladon The opooia Characters— Morphological hlgntcAnalysis— Phylogenetic Homalomena Homalomena EAF B LADE subgenus Breni ope)i nywal supported weakly only is complex) (Borneensis r ae pnoealifoecneshapes inflorescence overall upon based are P p r01fo h atcato Peninsular of coast east the from Ar3051 sp. OSTERIOR Homalomena Homalomena Pteromischum .vivens H. Homalomena L Homalomena OBES .expedita H. h uegopboundaries supergroup The p r05(etSrwk is Sarawak) (West Ar3065 sp. ssprtdit w clades: two into separated is Pseirlb definition lobe —Posterior Homalomena hc a ntal ten- initially was which , M 1,P .6 (clade 0.86) PP 61%, (ML Homalomena .philippinensis H. es aoe al. et Mayo sensu Homalomena. M 9,P 0.85). PP 79%, (ML aainvestigated. taxa .curvata H. r presented are < Homalomena Philodendron Homalomena 0,value 50%, (indicated .Thisis Poste- basal , nei h oaoeaadCaacao G.Spathe and SG SGs. Punctulata SG, Chamaecladon Neotropical Cyrtocladon in and present Homalomena is constriction the in once ptecntito speimrhcfor plesiomorphic is constriction Spathe hr r w bevtoswrhntn.Frt he markers three First, R peaks, noting. (at worth common observations For two and are S1). there selected Table data, were (supplementary markers scored chemical chro- Twelve the Asian supergroups. analyzed to unique or we common peaks chromatograms, for matograms LC from retrieved oeie iutdaoetefriezns(e.g. zones fertile the above situated sometimes ikdt comdtn ifrn ie fpollinators of sizes possibly is different obs.). pers. it Hoe, accommodating C. (Y. investigation; to further shorter linked needs is state spathe lower character in the spathe upper of the length than the complexes SG) at Borneensis (Cyrtocladon and inflorescences Giamensis the from for taken Except anthesis. are comparisons length spathe tmn;foeswt orsaesaeplesiomorphic are flower. stamens staminate per stamens four with two in by or flowers four obscured either has stamens; is flower staminate differentiation Each close-packing). the occasionally (although in flowers omn hmodlpaewe iwdfo above. expanded all from at visible. clearly not viewed are is anthers when SG the anthers, and Chamaecladon the plate in connective overtops rhomboidal The and a expanded is forming Chamaecladon connective fly-pollinated the known) SG, (where the of species iga h on orsodn otejnto ewe the between (e.g. junction spadix the the of zones to pistillate corresponding and staminate point the at of ring genera many in occurring spathe itllnt)ta h soitdpsisweesi the slightly in or whereas equal pistils they pistil. SGs the the associated exceed Cyrtocladon half the and exceeding than Homalomena (seldom length) shorter pistil much are staminodes ihrwt rwtotankdsie ycnrs h spadix the contrast By stipe. peduncle; Neotropical naked the a of without onto or obliquely) with (not either directly inserted spadix A S tmndsaeasn nteHmlmn G Punctulata genus SG, with Homalomena the the associated and for in SG, more, absent plesiomorphic are rarely is Staminodes staminode, pistil) (one each zone late ek nti einidctdta l ltdpasare peaks eluted all that indicated region this the in of except absorption UV-visible peaks samples of Analysis all chromatogram). for from chromatograms min expedita 6.50 H. the and min 5.90 in between peaks observed of 3). region (Fig. a species) also several is in There detected not (but analyzed groups pdxisrini hrfr naoopi hrce state character apomorphic Neotropical an the for therefore is insertion spadix iso h hmeldnS)adtePntlt G(includ- SG complexes). Havilandii Punctulata and the Insignis and ing SG) Chamaecladon the of cies TAMINODES SSOCIATED .L 3. .S 4. .S 8. dniiaino Markers— of Identification .S 7. .H 6. .S 5. .S 2. Homalomena t .5 .9 n .0mn r eetbei l h super- the all in detectable are min) 2.90 and 2.69, 2.55, PADIX PATHE HAPE TAMEN TAMINODES OWER EIGHT .vivens H. Homalomena Hmlmn G sesplmnaydt i.S2 Fig. data supplementary (see SG) (Homalomena —h rsneo tmndsaogtepistil- the among staminodes of presence )—The / O P Homalomena UPPER O I F ISTIL C NSERTION N F C ONSTRICTION I Homalomena UMBER ONNECTIVE NTERPISTILLAR ihteCaacao Ghvn two having SG Chamaecladon the with I h hmeldnS,interpistillar SG, Chamaecladon the —In A CroldnSG). (Cyrtocladon S MONG Homalomena. PATHE AlAsian —All r lasidvdal differentiated individually always are . P Homalomena T ER Acntitdprino the of portion constricted —A P L HE ER ENGTH sisre biul.Oblique obliquely. inserted is S S P S TAMINATE TAMINODE ISTILLATE rmteceia profiles chemical the From TAMINATE R ATIO Araceae h infcnei this in significance The . Homalomena Homalomena Telower/upper —The I Z F N F ONE LOWER LOWER eeal occur- generally ; C OMPARISON (I Cryptocoryne Homalomena Homalomena Ecp for —Except NTERPISTILLAR —Staminate Homalomena aethe have n lost and W ITH ). ), . Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. n ge l 21)aeidctda lc n ht oe.Cmlxso h Asian the of values Complexes (bootstrap branch boxes. the Neotropical white above and the shown black are of values as Erythropus indicated and are Crinipes (2011) and al. et Ng and RAgns ld ...represents Pteromischum O.W.H. Clade genes. rRNA 03 OGE L:PYOEYO SA OAOEA(RCA)593 (ARACEAE) HOMALOMENA ASIAN OF PHYLOGENY AL.: ET WONG 2013] Fig. .Mxmmlklho hlgn f8 aa( taxa 89 of phylogeny likelihood Maximum 1. . = P.P , Philodendron subgenus Homalomena Philodendron Homalomena pce rmteOdWrdtois .. e ol rpc,P t.= Pte. P. tropics, World New = N.W.H tropics, World Old the from species < Homalomena 0 r o shown). not are 50% n ..= P.M. and , r niae nbaklns otta mxmmlklho)adpseirprobability posterior and likelihood) (maximum Bootstrap lines. black in indicated are , Philodendron Philodendron n ugop)bsdo T1 T2 n h .Srgo ftenuclear the of region 5.8S the and ITS2, ITS1, on based outgroups) and subgenus Homalomena Meconostigma es oc n og(08 n ge l (2011), al. et Ng and (2008) Wong and Boyce sensu uegop es oc n og(2008), Wong and Boyce sensu supergroups . Philodendron subgenus Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. agdfo –.3adR,fo –.7 w markers Two 0–0.57. from RI, values CI and the 0–0.33 as homoplasies from high ranged showed excluding markers min) 4.01 these and 3.92, 3.88, 3.81, tmnds,(i egto nepsilrsaioei oprsnwt soitdpsi,(i)Sae ubrprsaiaefoe,ad(ii Sh (Interpis (viii) zone and pistillate flower, staminate characters. the morphological per among each number for Staminodes shown Stamen are (v) (vii) (RI) insertion, pistil, index retention associated Spadix and with (CI) (iv) comparison index ratio, Consistency in stamen. length staminode per connective spathe interpistillar of of Lower/upper Height (iii) (vi) constriction, staminodes), Spathe (ii) lobes, aiaa 2,40 n 6 m(oaah ta.20) and 2006), al. analysis. character-mapping et the (Kobayashi in further nm studied absorption 660 not hence and UV-visible 410, 226, with at maxima structures chlorophyll-type of 38 [Volume BOTANY SYSTEMATIC 594 eefudt epeeti oto h ebr (R members the markers of most five unique in SG, present as Cyrtocladon be identified the to markers found In of were supergroup. number each a to are there taxa, te hntemrescmo cosmany across common markers the than Other Fig. .Egtmrhlgclcaatr piie noasmaie oeua hlgn rmFg for 1 Fig. from phylogeny molecular summarized a onto optimized characters morphological Eight 2. .vivens H. Homalomena However, . t 3.74, I=08)wr on ob hrdb h Chamaecladon, (R species the markers Neotropical by For shared SGs. be (only Homalomena to found and were Punctulata 0.80) = RI (R cs(uknhm21) twsosre htms fthe of most that Prod- observed Natural was of It 2011). Dictionary cross-referenced (Buckingham database: data ucts commercial MS a resultant the with spec- with mass by (MS) analyzed were trometry samples the all markers, twelve t dniyo Markers— of Identity .0mn I=05,R .7 n .0mn I=0.50, = CI min, 3.60 and 0.67, = RI 0.50, = CI min, 2.80 .wallisii H. t .5ad35 i)wr identified. were min) 3.54 and 3.25 a nlzdi h hmclpoiig,two profiling), chemical the in analyzed was ocaatrz h dniyo the of identity the characterize To Homalomena i efbaeposterior blade Leaf (i) . tillar ape Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 03 OGE L:PYOEYO SA OAOEA(RCA)595 (ARACEAE) HOMALOMENA ASIAN OF PHYLOGENY AL.: ET WONG 2013] x)ad(i)aeuiu oteNeotropical and the to SG unique Homalomena are SG, (xii) and Chamaecladon (xi) in detected were (ix) Homalomena Fig. .Tev hmclmresietfe rmLM rfln,otmzdot umrzdmlclrpyoeysoni i.1for 1 Fig. in shown phylogeny molecular summarized a onto optimized profiling, LCMS from identified markers chemical Twelve 3. hmclmres i o(i)wr eetdi lotaltetx nlzd i)t vi)aeuiu oteCroldnsprru (SG); supergroup Cyrtocladon the to unique are (viii) to (iv) analyzed; taxa the all almost in detected were (iii) to (i) markers: Chemical . Homalomena. ossec ne C)adrtninidx(I r hw o ahceia characters. chemical each for shown are (RI) index retention and (CI) index Consistency .punctulata H. x eedtce nCaacao,Hmlmn n ucuaaSGs; Punctulata and Homalomena Chamaecladon, in detected were (x) ; Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 9 YTMTCBTN Vlm 38 [Volume BOTANY SYSTEMATIC 596 to that suspected is It petioles. Andre bevdt evstdol by only SGs Cyrtocladon not visited and Homalomena connective whereas be Drosophilidae), the are SG to with Chamaecladon the observed stamens of Species thecae. two the overtopping with staminate and each pistil, interpistillar associated having flowers the by morphologically than SGs shorter two is other staminodes SG the to Among Chamaecladon dissimilar the the SGs. most of SGs, Punctulata species three most + these by Homalomena shared + chem- and Two Chamaecladon unique be SG. are Cyrtocladon to of markers detected species ical the were of most markers to chemical unique Five the in zone. staminodes leaf of pistillate on supported presence lobes is and SG. posterior spathe, and of constricted SGs, blades, presence Punctulata other having others, the into among to by, sister complexes is SG Insignis Cyrtocladon the and of removal the Havilandii after monophyletic is SG Punctulata Cyrtocladon SGs. and Homalomena, Chamaecladon, Cyrtocladon, ihla ldso og lbosptoe ut ifrn from of different illustration quite plant) petioles (Type original glabrous the long, plant on solitary blades a leaf is represents with specimen herbarium probably the and novelty; misidentified, taxonomic is a (2008) al. et of Gauthier specimen The leaves. whereas, leaves, and indumentose and staminodes and spe congruity. the morphological with 1), (Fig. Erythropus characters. free morphological often and Neotropical phylogeny and the insertion with straight congruent Asian a in insertion to stipitate compared as Neotropical junction of spadix the with insertion Homalomena spadix 2): Fig. to (refer Ma 25 Homalomena least at (Nauheim recent, Oligocene divergence relatively the estimated during is groups The two Asian). the these is for between type retained (the name species generic Asian the with Asian species, strongly of Neotropical here separation presented generic data support associated the and phylogeny nteCroldnS eeas on ob rsn nthe in present be to found also were outgroup, to SG unique Cyrtocladon be the in not plant may many here plant in Homalomena identified of present markers group be the to a families, secondary known are plant metabolites flavonoids of secondary class as Harborne useful However, by most metabolites. qualified the probably been systematic as have the (1973) from flavonoids (Williams that view, noteworthy studied of further species point is It Araceae 1981). al. of et possible, (82%) flavonoids a majority in of found high were types C-glycoflavones many or C-glycosides the flavones occur- the Of metabolites Araceae. common in the among ring by be pigments records to anthocyanin flavonoids chemotaxonomy reported and which the (1981), 2005). al. with al. et et agreement Williams (Ulubelen in nm is 240–280 the and This nm at peaks 300–380 absorption of major range two of exhibit characteristic which profiles flavonoids, UV-visible have markers identified Homalomena oaoea urnl oyhltcAssemblage— Polyphyletic a Currently Homalomena, .kvistii H. .wallisii H. ´ .panamamensis H. 83,wihi lmigpatwt hr,pilose short, with plant clumping a is which 1873), .nervosa S. Croat. ned oeo h akr htw observed we that markers the of some Indeed, . ld sspotdb n opooia character morphological one by supported is clade biul netdo h oe spathe/peduncle lower the on inserted obliquely Homalomena akitritla tmndsadhv glabrous have and staminodes interpistillar lack es tit opie orsupergroups: four comprises stricto sensu . hr h rsneo interpistillar of presence the share oaoeapicturata Homalomena oaoeacrinipes Homalomena Homalomena omtocae:Ciie and Crinipes clades: two form .‘picturata’ H. re l 02) h Neotropical The 2012b). al. et er isi h ldsdisplaying clades the in cies Colocasiomyia .picturata H. hmclmresare markers Chemical . Homalomena ape eei closer is here sampled , nlddfrom included .wendlandii H. .erythropus H. Lne and (Linden rmthe from (Diptera: The efta a,i oeseis edel odt (Grayum cordate (usually) deeply be a species, article: some expanded in per sheathless may, virtually that produced a leaf and leaves cataphyll two conspicuous only with n infcn iiaiisto similarities significant and G hyaesprtdb h bec rol eyweak very a only or absence the by separated species Cyrtocladon are to Although they similar SG, vegetatively needed. look SG are Homalomena of mechanisms investigations pollination further of obs.); pers. (P.C.Boyce, obser- inflorescences and taxa, of these vations in mechanism pollination the in alternative zone pollina- and pistillate SG the destructive among Punctulata are staminodes which of beetles, Absence tors. from protection anthers provides expanded the probably The for SG 2011a). al. Cyrtocladon et in Hoe beetle connective 2010; probably al. most et are (Tung rewards, pollinated food as by functioning as ingly well Rutelinae, (Scarabaeidae: as Colocasiomyia beetles Staphylinidae), of Nitidulidae, variety Chrysomelidae, a by visited are and iiaiiswt subgenera with similarities aresisterto the by and spathe, the of movements. closing spadix of and absence gaping anthesis at simple spathe, upper the and lower the between constriction acoaet litcl vt,o cainlysubcordate occasionally of subgenera or other ovate, Both and elliptical, blades. petioles to sheathed extensively and lanceolate while have inconspicuous), article, leaves highly least expanded stem at all per (or leaves absent many are cataphylls to several with vegetative adult shoots, in growth sympodial anisophyllous exhibits subgenus predated. be the of to in observed that tissues chemical note vegetative to plant’s interesting the field is It of herbivores. part against form arsenal irritating doubtless much-reported the that in properties involved lacking be to SG suspected and Cyrtocladon in species only the staminodes. interpistillar being in tive Elopium Elopium xmnto fseisof ( species of genus examination lianescent predominantly and ( mesophytes ( undertaken. sampling, are comprehensive changes more of much especially and changes, nomic of rest the to ecmie with combined be of rest Neotropical the from of separation the Neotropical and of rest the to sister are eeacnb ananda eaaetaxa, separate as maintained be can genera Elopium subgenus ‘ of Neotropical one but all of remainder Pteromischum aooi Implications— Taxonomic origin phenolic of were detected compounds chemical All hl tmyse mluil htacaeo lianas of clade a that implausible seem may it While Neotropical Meconostigma novn oigall moving .Thiswillresultin involving , Sht 85 sagnrcsnnm lentvl,the Alternatively, synonym. generic a as 1865) (Schott Pteromischum .expedita H. Philodendron Philodendron n ae ntedmg osaioe seem- staminodes to damage the on based and , Pteromischum Adelonema Pteromischum smr lsl eae oacaeo terrestrial of clade a to related closely more is ) Philodendron ’Homalomena’ Philodendron Homalomena Philodendron neg ihlossmoilgrowth, sympodial diphyllous undergo ) .expedita H. Philodendron r htonly that are hnt nohriemonophyletic otherwise an to than ) sagns( genus a as subgenus Homalomena’ hs r aetysgiiattaxo- significant patently are These . hswudrqierecombining require would This . srqie eoeteeformal these before required is , into Pteromischum eea pin exist: options Several . Adelonema Homalomena 1 ecie pce)myeither may species) described (10 Philodendron Neotropical sa es swl upre as supported well as least at is es ao rwith or lato, sensu Hmlmn G mle an implies SG) (Homalomena ’Homalomena’ oaoeavivens Homalomena Adelonema h eaaino h Asian the of separation The . Pteromischum Pteromischum Philodendron dlnm.Pteromischum Adelonema. Colocasiomyia Adelonema pce n l species all and species + Philodendron Elopium eeams never almost were eelsrkn dis- striking reveal Homalomena and Sht 80 with 1860) (Schott + Adelonema Meconostigma )sistertothe ,andtogether ( Pteromischum pce into species Philodendron Philodendron en sister being tedthe attend sdistinc- is ,closer ), species and , Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ale,M 2011. M. Carlsen, arr,L . .A aaa,M .Cae .J ao .Bge,and Bogner, J. Mayo, J. S. Chase, W. M. Salazar, A. G. I., L. Cabrera, 2011. J. Buckingham, oc,P . .Y og .L o,A .J ig .E o,I .Oi and Ooi, H. I. Low, E. S. Ting, J. P. A. Low, L. S. Wong, Y. S. C., P. Boyce, oc,P .adS .Wn.20.Suiso oaoeee(Araceae) Homalomeneae on Studies 2008. Wong. Y. S. and C. P. Boyce, oc,P .adH i 2010. Li. tribe H. and Araceae C. of P. revision Boyce, taxonomic A 2001. Hay. A. and C. P. U Boyce, The 2011. Croat. B. T. and C. P. Boyce, oc,P .20b h genus The 2001b. C. P. Boyce, oc,P .20c h genus The 2000c. C. P. Boyce, oc,P .20a h genus The 2001a. C. P. Boyce, genus The 2000b. C. P. Boyce, genus The 2000a. C. P. Boyce, Homalomeneae on Studies 2010. Barabe Boyce. C. P. and identification. S. model statistical Baharuddin, the at look new A 1974. H. Akaike, grate- are Centre Biodiversity Sarawak Sarawak the the acknowledged. of fully and support & and Department VI)-41 collaboration The (Jld. Forestry 27/2011. NCCD.907.4.4 No. No. Permit Permit Park Research Sarawak 09–173 Department using conducted No: Forestry recently Grant most was Research Fieldwork RG/PHA/AS_C. COMSTECH–TWAS: and 05-01-09-SF0006 onr .J .16.TeMlynfoa p 12.in 21–24. Pp. flora. Malayan The 1960. H. J. E. Corner, Kuching: Kelabit. the and Iban the of Ethnobotany 2002. H. Christensen, 03 OGE L:PYOEYO SA OAOEA(RCA)597 to assigned currently Adelonema Neotropical of Asian species least published the very the of at cataphylls and the Homalomena prophylls (ARACEAE) to HOMALOMENA contrast developed ASIAN (in OF PHYLOGENY cataphylls highly AL.: inconspicuous ET article, WONG leaves stem spathe. many per to several with the growth sympodial spadixanisophyllous the surpassing with and free, distally ultimately and persis- involute sheath often the and with tent, petioles characterised geniculate” further “apically having (1860) Schott 1996). 2013] wisfrhrivsiaint nld oeseisof species more include to investigation Adelonema further awaits Acknowledgments. Anthurium DNA. plastid Botany noncoding of and Journal American coding from inferred (Araceae) weeds .Da P. .K g 00 rca fBorneo. of Araceae 2010. Ng. K. K. fBre :Fu e pce n pclto nifra species informal on speculation and species Sarawak. in new groups Four Bogner I: J. Borneo Hetterscheid, of A. L. W. Murata, J. 23:17–18. Jacobsen. Boyce, N. C. and P. Zhu, H. G. n h rpclWsenPacific. Western Genera. tropical the and Aroid in ( Potheae species of number estimated http://www.aroid.org/genera/111109uberlist.pdf. and published Pacific. western tropical the and Gardens’ Australia Guinea, New in Monstereae) oseee ntePhilippines. the in Monstereae) oseee nBorneo. in Monstereae) Indochina. and Thailand of archipelago. Indonesian western and southern Gardens’ the in Monstereae) phylogeny and flowers (Araceae). bisexual the atypical in of development between Perak from I: Malaysia Peninsular of (Araceae) Control Automatic on Transactions fcneayadbcneaycnrs fbiology of Malaya. congress of University bicentenary Singapore: and and centenary of Denmark NEPCon Malaysia, Aarhus. of Sarawak, University Department Forest fossils and phology ´ . .Bueu .Frs,adC ari.20.Tecorrelation The 2002. Lacroix. C. and Forest, F. Bruneau, A. D., , ´ ia 08 hlgntcrltosiso risadduck- and aroids of relationships Phylogenetic 2008. vila. h ttsadgnrcnmnltr ftespecies the of nomenclature generic and status The . n etrrpeetto of representation better a and . . ultnSingaporeBulletin ultnSingaporeBulletin ohs Pothoidium Pothos, ,adetnieysete eils nconclusion, In petioles. sheathed extensively and ), caPyoaooiaGeobotanica Phytotaxonomica Acta cot(rca) nertn oeua hlgntc,mor- phylogenetics, molecular integrating (Araceae), Schott nesadn h rgnadrpddvriiaino h genus the of diversification rapid and origin the Understanding itoayo aua products natural of Dictionary h .dsetto.S oi:Uiest fMissouri. of University Louis: St dissertation. D. Ph. . 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E, S. 110.16 Temple, N; 01.19 Village, Mariamman Giam Padawan, Kuching, Sarawak: E, Maha 110.13 Matang, N; 01.35 Kuching, Ar2574 Sarawak: Wong MALAYSIA. Y. E, S. 110.20 N; 01.20 Village, Sikog Padawan, E, Kuching, Sarawak: 112.03 N; Ar2389 01.12 al. Ai, Batang et Antu, Lubok Aman, Sri Sarawak: * MBG52988, DQ866880. (MO), Temple, Mariamman Homalomena E, Maha 110.13 Matang, N; 01.35 Kuching, Sarawak: MALAYSIA. E, 112.03 N; JX076784. 01.12 (SAR), Ai, Batang Antu, Lubok Aman, r mn uo nu aagAi, Batang Antu, Ar3004 Lubok Aman, Sri sengkenyang * AASA aaa:SiAa,LbkAt,Btn i 11 ;112.03 N; 01.12 Ai, Batang Antu, E, Lubok Aman, Sri Sarawak: MALAYSIA. E, MBG90199, AASA aaa:SiAa,LbkAt,Btn i 11 N; 01.12 Ai, Batang Antu, Lubok E, Aman, 112.03 Sri Sarawak: E, MALAYSIA. 112.03 E, N; 112.03 N; 01.12 01.12 Ai, Batang Ar2362 Antu, Lubok Aman, Sri N; 00.00 Sekadau, sengkenyang Kalimantan: West E, 111.04 Kalimantan: N; INDONESIA. 01.12 Ai, Griff. Batang Antu, Lubok E, 112.03 Aman, Sri Sarawak: MALAYSIA. 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MBG79249, E, 110.13 N; Ar3057 01.35 Temple, Mariamman Maha Matang, Kuching, oaoeajosefii Homalomena oaoeasengkenyang Homalomena oaoeahumilis Homalomena oaoeaatrox Homalomena .C oc ta.A 2414 Ar al. et Boyce C. P. oaoeaasmae Homalomena .C oc ta.Ar2361 al. et Boyce C. P. SR,JX076778.(SAR), SR,JQ955574. (SAR), SR,J068;Srwk r mn uo nu aagAi, Batang Antu, Lubok Aman, Sri Sarawak: JX076787; (SAR), nl AASA aag eatt ruWllf ete 35 N; 03.50 Centre, Wildlife Krau Jerantut, Pahang: MALAYSIA. Engl. nl– MBG81956c, Engl.–, SR,JX076790. (SAR), .C oc .Y ogAr2532 Wong Y. S. & Boyce C. P. .C oc ta.Ar2424 al. et Boyce C. P. .C oc ta.Ar3052 al. et Boyce C. P. oaoeaclandestina Homalomena oaoeadebilicrista Homalomena .C oc ta.Ar2388 al. et Boyce C. P. oaoeahanneae Homalomena SR,J067.* JX076779. (SAR), ra 79249 Croat ra 52988 Croat gKa iw&JpmA ia Ar3005 Tisai Ak Jipom & Kiaw Kiaw Ng ra 90199 Croat SR,JQ955571. (SAR), ..oc,SYWn aiudn AASA Sarawak: MALAYSIA. Fasihuddin. & S.Y.Wong P.C.Boyce, ..oc,SYWn aiudn AASA Sarawak: MALAYSIA. Fasihuddin. & S.Y.Wong P.C.Boyce, aff. .C oc eadA ia Ar2451 Kisai Ak Jeland & Boyce C. P. oaoeamatangae Homalomena nl ITA.Nn ihPoic:CcPhuong Cuc Province: Binh Ninh VIETNAM. Engl. oaoeagiamensis Homalomena panamense SR,J068.Srwk uhn,Mtn National Matang Kuching, Sarawak: JX076782. (SAR), .C oc eadA ia Ar227 Kisai Ak Jeland & Boyce C. P. .C oc eadA ia Ar230 Kisai Ak Jeland & Boyce C. P. oaoeainsignis Homalomena oaoeaphilippinensis Homalomena ..Bye ..Wn aiudn MALAYSIA. Fasihuddin. & Wong S.Y. Boyce, P.C. SR,J060,J060.* JX076804. JX076803, (SAR), SR,J957.* JQ955573. (SAR), ..oc ..og AASA aaa:Sri Sarawak: MALAYSIA. S.Y.Wong. & P.C.Boyce SR,JQ955572. (SAR), SR,J992.* JQ929129. (SAR), .C oc ta.Ar2387 al. et Boyce C. P. M) DQ866881. (MO), O,D867.* DQ866879. MO), oaoea‘gastrofructa’ Homalomena Jc)Hook. (Jack) oaoeaerythropus Homalomena aaudn&P .Bye AASA Perak: MALAYSIA. Boyce. C. P. & Baharuddin SR,JX076773. (SAR), il AASA aaa:Kcig Bako Kuching, Sarawak: MALAYSIA. Ridl. M) Q682 * DQ866882. (MO), .C oc ta.Ar2111 al. et Boyce C. P. ra 77907 Croat oaoeahanneae Homalomena Homalomena SR,JX076772. (SAR), ra 81956 Croat ra acl.,MBG90162, Marcell.–, & Croat oaoea‘baangongensis’ Homalomena .C oc ta.Ar2360 al. et Boyce C. P. ..oc,SYWn Fasihuddin. & S.Y.Wong P.C.Boyce, aaudnAr2597 Baharuddin SR,JX076785. (SAR), ..oc,SYWn Fasihuddin. & S.Y.Wong P.C.Boyce, f AASA aaa:SiAman, Sri Sarawak: MALAYSIA. . ..oc,SYWn Fasihuddin. & S.Y.Wong P.C.Boyce, SR,J067;Phn:Jerantut, Pahang: JX076776; (SAR), gKa iw&JpmA Tisai Ak Jipom & Kiaw Kiaw Ng ..o.MLYI.Sarawak: MALAYSIA. Y.C.Hoe. .C oc ta.Ar2385 al. et Boyce C. P. oaoeahanneae Homalomena SR,JX076789. (SAR), ..ug ..og&P.C.Boyce. & S.Y.Wong L.S.Tung, ..o,SYWn P.C.Boyce. & S.Y.Wong Y.C.Hoe, M) DQ866877. (MO), oaoeapicturata Homalomena SR,J068.* JX076786. (SAR), oaoeaexpedita Homalomena .C oc ta.Ar3053 al. et Boyce C. P. oaoeaborneensis Homalomena oaoeaborneensis Homalomena M) DQ866878. (MO), SR,JX076775. (SAR), ..r AASA Sarawak: MALAYSIA. N.E.Br. .Byee l Ar2371 al. et Boyce C. P oaoeaclandestina Homalomena p AASA Sarawak: MALAYSIA, sp. oaoea‘chartacea’ Homalomena oaoeapunctulata Homalomena nl ssp. Engl. .C oc ta.Ar2380 al. et Boyce C. P. .MALAYSIA.Sarawak: oaoeahanneae Homalomena ..oc,SYWn & S.Y.Wong P.C.Boyce, .C oc l Ar2575 al. & Boyce C. P. oaoearostrata Homalomena nl PHILIPPINES. Engl. SR,JX076771. (SAR), SR,JX076788. (SAR), .C oc tal. et Boyce C. P. SR,JX076783. (SAR), SR,JX076780. (SAR), SR,JQ955578; (SAR), SR,JQ955577. (SAR), SR,JX076781. (SAR), .C oc tal. et Boyce C. P. SR,JX076801, (SAR), gKa iw& Kiaw Kiaw Ng allenii MALAYSIA. .  Homalomena Homalomena Homalomena Homalomena Homalomena .C oc & Boyce C. P. .C oc & Boyce C. P. 1 ;112.03 N; .12 ra 90162 Croat .C Boyce C. P. .C Boyce C. P. P.C.Boyce, .C Boyce C. P. .C Hoe C. Y. .a & A.Hay Regel.–, Croat.–, (SAR), (SAR), (SAR), .C. P. Ridl. Ridl. P.C. . Delivered by Publishing Technology to: Sin Yeng Wong IP: 161.142.87.2 on: Sun, 22 Sep 2013 23:59:17 Copyright (c) American Society for Plant Taxonomists. All rights reserved. DQ866895. 599 (ARACEAE) HOMALOMENA ASIAN OF PHYLOGENY AL.: ET WONG JX076791. Ai, Batang Antu, Lubok E, 102.20 JX076792. N; 02.24 (SAR), Sedana, Bukit Simpan Hutan Ar3065 2013] nu aagAi, Lubok Aman, Batang Sri Ar2363 Sarawak: Antu, MALAYSIA. al. Ai, Fasihuddin. & Batang et S.Y.Wong Antu, Boyce, Boyce Lubok C. Aman, P. Sri * Sarawak: MALAYSIA. Fasihuddin. JX076809. Ai, (SAR), *Batang Antu, Antu, Lubok Lubok Aman, Aman, E, 112.03 Sri JX076808. N; Sarawak: 01.12 E, MALAYSIA. Ai, 112.03 Batang Fasihuddin. N; & 01.12 Wong Ai, Batang Ar2390 Antu, Lubok Aman, SR,J069.* E, JX076793. 103.21 (SAR), N; 02.37 Centre, Wildlife M) DQ866897. (MO), DQ866896. (M0), MBG81932a, Nursery, Landscaping Borneo ex cultivated * JX076797. hldnrngoeldii Philodendron fragrantissimum MBG38218, Grayum.–, DQ866887. JBM145–2003, JBM1518–2003,–, Schott.–, 38 Gauthier 2 wendlandii Homalomena DQ866883. glaziovii M) DQ866884. (MT), SR,JX076794. (SAR), SR,J069.* JX076795. (SAR), oaoeavagans Homalomena Hmlmn ‘stella’ *Homalomena oaoeawallisii Homalomena hldnrnangustisectum Philodendron hldnrngrandipes Philodendron Hook. M) DQ866885. (MT), ra 81932 Croat atir1 Gauthier ut.,FG, Kunth.–, hldnrnheleniae Philodendron Homalomena Homalomena hldnrnhylaeae Philodendron gKa iw&JpmA ia Ar3006 Tisai Ak Jipom & Kiaw Kiaw Ng hldnrnanisotomum Philodendron f. oaoeavivens Homalomena ,BM7014–1998, –, ars.,JBM1699–1996, Barroso.–, Homalomena M) DQ866890. (MT), ra 38212 Croat gKa iw&JpmA ia Ar3033 Tisai Ak Jipom & Kiaw Kiaw Ng oaoeasymplocarpifolia Homalomena M) DQ866886. (MO), SR,J069.* JX076796. (SAR), cot– MBG85114a, Schott.–, hldnrnbarrosoanum Philodendron .C oc ta.Ar2411 al. et Boyce C. P. Barabe p AASA aag eatt Krau Jerantut, Pahang: MALAYSIA. sp. AASA aaa:Srwk Sri Sarawak, Sarawak: MALAYSIA. . gKa iw&JpmA ia Ar3011 Tisai Ak Jipom & Kiaw Kiaw Ng ee.MLYI.Srwk Kuching, Sarawak: MALAYSIA. Regel. p AASA aaa:SiAman, Sri Sarawak: MALAYSIA. sp. ..oc.MLYI.Srwk Sri Sarawak: MALAYSIA. P.C.Boyce. .C oc gKa iwAr3051 Kiaw Kiaw Ng & Boyce C. P. ´ rue– MBG79244, Krause.–, 77 p AASAMlk:Machap, Melaka: MALAYSIA sp. M) DQ866892. (MO), hldnrnecordatum Philodendron nl– B 2801–1950, JBM Engl.–, ra,,MBG83278, Croat,–, M) DQ866893. (MT), hldnrnfindens Philodendron ..utn.,MBG84578a, G.S.Bunting.–, hldnrnbrevispathum Philodendron atir26 Gauthier ..oc,SYWn & S.Y.Wong P.C.Boyce, oaoeavivens Homalomena Ar3603 cot– JBM2803–1950, Schott.–, .C oc ta.Ar3047 al. et Boyce C. P. ra 85114 Croat atir27 Gauthier .C oc tal. et Boyce C. P. SR,JX076798. (SAR), SR,JX076807, (SAR), ..oc,S.Y. P.C.Boyce, DQ866894. , G.S.Bunting.–, Philodendron Philodendron ra 79244 Croat ra 83278 Croat ra & Croat Gauthier Schott. (SAR), (SAR), (MO), (MT), P.C. atir31 Gauthier 75 1996, 6 Chouteau DQ866903. DQ866902. atir37 Gauthier M) DQ866901. (MT), 84578 Croat rnsolimoense dron MBG55184c, DQ866904. (MT), ok– B 97–100, smithii MBG Hook.–, Garden, Botanical Penang DQ866909. sagittifolium Garden, Botanical dendron Penang ex cultivated DQ866908. (MT), DQ866907. pterotum JBM2043–1997, Kunth.–, ta.14 al. et dron okr ..og&PCBye RNHGUIANA. FRENCH P.C.Boyce. & S.Y.Wong Jonker) Boos.–, sodiroi DQ866916. dendron Philodendron rnsquamiferum dron a,MutBd,0.3N 1.7E, 110.07 N; JX076800. 01.23 (SAR), Bidi, Mount Bau, * JN544445. M) DQ866899. (MT), atir33 Gauthier p ( sp. MBG71897 nl– MBG64524, Engl.–, ot– BM7163–1995, Hort.–, K) DQ866918. (KW), p,,JBM1659–1953, sp.,–, .oh&Agsi.,JBM1840–1955, Augustin.–, & K.Koch cimtgotsnervosa Schismatoglottis M) DQ866888. (MT), hldnrnsurinamense Philodendron pteromischa hldnrnradiatum Philodendron hldnrnlundii Philodendron hldnrnmamei Philodendron hldnrnselloum Philodendron M) DQ866898. (MO), M) DQ866900. (MT), M) DQ866919. (MT), ra 55184 Croat p– JBM1130–1952, sp.–, ..mt.,FG, A.C.Smith.–, hldnrnlongistilum Philodendron DQ866920. , M) DQ866891. (MT), hldnrnradiatum Philodendron op.&Ed.,JBM7009–1998, Endl.–, & Poepp. hldnrnmartianum Philodendron .,MBG84914, ).–, hldnrnlindenii Philodendron ib.,JBM3402–1983, Liebm.–, ra 97–100 Croat atir44 Gauthier hldnrnundulatum Philodendron g2 Ng ra 64524 Croat M) DQ866905. (MO), hldnrnornatum Philodendron hldnrninsigne Philodendron hlnto americanum Philonotion hldnrnlinnaei Philodendron hldnrnxanadu Philodendron atir19 Gauthier atir36 Gauthier SR,JX089318. (SAR), am MBG82932, Warm. nr.,BM7224–1992, Andre.–, Barabe .oh AASA utvtdat cultivated MALAYSIA. K.Koch. atir20 Gauthier il AASA aaa:Kuching, Sarawak: MALAYSIA. Ridl. cot– JBM2802–1950, Schott.–, hldnrnpanamense Philodendron .C oc eadA ia Ar944 Kisai Ak Jeland & Boyce C. P. Mq xSht)Eg.,FG, Engl.–, Schott) ex (Miq. M) DQ866911. (MO), M) DQ866906. (MT), ra 84914 Croat ´ M) DQ866912. (MO), g1 Ng 60 cot AASA Penang: MALAYSIA. Schott. rue– MBG-11–17–78,–, Krause.–, M) DQ866917. (MT), M) DQ866913. (MT), M) DQ866914. (MT), als– JBM7064–1998, Wallis.–, M) DQ866910. (MT), SR,JX076799. (SAR), nl– JBM2424–1946, Engl.–, hldnrnpedatum Philodendron hldnrnserpens Philodendron nl– JBM1930–52, Engl.–, cot– FG, Schott.–, atir46 Gauthier cot– JBM1511– Schott.–, Kunth.–, atir42 Gauthier onr2911 Bogner ra 82932 Croat ra,My J. & Mayo Croat, M) DQ866915. (MO), atir45 Gauthier AME okr& Jonker (A.M.E. Philodendron Philodendron Philodendron atir39 Gauthier atir6 Gauthier Philoden- Philoden- Philoden- Krause.–, Barabe Philo- Philo- (SAR), Barabe (MO), (MT), (MT), (MT), Haig ´ 76 ´